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3.1 Extensions to Mendel for Single-Gene Inheritance    47


                       Incomplete dominance: The F 1  hybrid resembles     with an enzymatic role in red pigment production. The
                                                                                        2
                       neither pure-breeding parent                        white allele (A ) does not give rise to a functional enzyme,
                                                                                            1
                       A cross between pure late-blooming and pure early-blooming   but the red allele (A ) does. Thus, in snapdragons, two red
                                                                                           1 1
                       pea plants results in an F 1  generation that blooms in be-  alleles per cell (A A ) produce a double dose of a red-
                       tween the two extremes. This is just one of many examples   producing enzyme, which generates enough pigment to
                                                                                                                         1 2
                       of incomplete dominance, in which the hybrid does not   make the flowers look fully red. In the heterozygote (A A ),
                       resemble either pure-breeding parent. F 1  hybrids that differ   one copy of the red allele per cell results in only enough
                       from both parents often express a phenotype that is inter-  pigment to make the flowers look pink. In the homozygote
                                                                                               2 2
                       mediate between those of the pure-breeding parents. Thus,   for the white allele (A A ), where there is no functional
                       with incomplete dominance, neither parental allele is dom-  enzyme and thus no red pigment, the flowers appear white.
                       inant or recessive to the other; both contribute to the F 1
                       phenotype. Mendel observed plants that bloomed midway
                       between two extremes when he cultivated various types of   Codominance: The F 1  hybrid exhibits traits
                       pure-breeding peas for his hybridization studies, but he did   of both parents
                       not pursue the implications. Blooming time was not one of   A cross between pure-breeding spotted lentils and pure-
                       the seven characteristics he chose to analyze in detail, al-  breeding dotted lentils produces heterozygotes that are both
                       most certainly because in peas, the time of bloom was not   spotted and dotted (Fig. 3.4a). These F 1  hybrids illustrate a
                       as clear-cut as seed shape or flower color.         second significant departure from complete dominance. The
                          In many plant species, flower color serves as a striking   progeny look like both parents, which means that neither
                       example of incomplete dominance. With the floret clusters   the spotted nor the dotted allele is dominant or recessive to
                       of snapdragons, for instance, a cross between pure-breeding   the other. Because both traits show up equally in the hetero-
                       red-flowered parents and pure-breeding white yields hy-  zygote’s phenotype, the alleles are termed  codominant.
                       brids with pink blossoms, as if a painter had mixed red and   Self-pollination of the spotted/dotted F 1  generation gener-
                       white pigments to get pink (Fig. 3.3a). If allowed to self-  ates F 2  progeny in the ratio of 1 spotted : 2 spotted/dotted : 1
                       pollinate, the F 1  pink-blooming plants produce F 2  progeny   dotted. The Mendelian 1:2:1 ratio among these F 2  progeny
                       bearing red, pink, and white flowers in a ratio of 1:2:1    establishes that the spotted and dotted traits are determined
                       (Fig. 3.3b). This is the familiar genotypic ratio of an ordi-  by alternative alleles of a single gene. Once again, because
                       nary single-gene F 1  self-cross. What is new is that because   the heterozygotes can be distinguished from both homozy-
                       the heterozygotes look unlike either homozygote, the phe-  gotes, the phenotypic and genotypic ratios coincide. 
                       notypic ratios are an exact reflection of the genotypic ratios.  In humans, some of the complex membrane-anchored
                          The simplest biochemical explanation for this type of   molecules that distinguish different types of red blood cells
                       incomplete dominance is that each allele of the gene under   exhibit codominance. For example, one gene (I) with
                                                                                        B
                                                                                  A
                       analysis specifies an alternative form of a protein molecule   alleles I  and I  controls the presence of a sugar polymer
                       Figure 3.3  Pink flowers are the result of incomplete dominance. (a) Color differences in these snapdragons reflect the activity of
                       one pair of alleles. (b) The F 1  hybrids from a cross of pure-breeding red and white strains of snapdragons have pink blossoms. Flower colors
                       in the F 2  appear in the ratio of 1 red : 2 pink : 1 white. This ratio signifies that the alleles of a single gene determine these three colors.
                       a: © Henry Hemming/Getty Images RF
                       (a)  Antirrhinum  majus (snapdragons)                (b)  A Punnett square for incomplete dominance


                                                                                                     1 1
                                                                                                               2 2
                                                                                     P              A A       A A
                                                                                     Gametes        A 1        A 2



                                                                                                     1 2
                                                                                                                1 2
                                                                                     F  (all identical)  A A   A A
                                                                                      1
                                                                                     F 2                     A 1  A 2
                                                                                                         A 1
                                                                                                             1 1
                                                                                                                  1 2
                                                                                                            A A  A A
                                                                                                         A 2
                                                                                                                  2 2
                                                                                                             1 2
                                                                                                            A A  A A
                                                                                                                     2 2
                                                                                                            1 2
                                                                                                   1 1
                                                                                                 1 A A  (red) : 2 A A  (pink) : 1 A A  (white)
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