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48     Chapter 3    Extensions to Mendel’s Laws


              Figure 3.4  In codominance, F 1  hybrids display the traits   Figure 3.2 summarizes the differences between com-
              of both parents. (a) A cross between pure-breeding spotted lentils   plete dominance, incomplete dominance, and codominance
              and pure-breeding dotted lentils produces heterozygotes that are   for phenotypes reflected in color variations. Determina-
              both spotted and dotted. Each genotype has its own corresponding   tions of dominance relationships depend on the phenotype
                                                  B
                                             A
              phenotype, so the F 2  ratio is 1:2:1. (b) The I  and I  blood group
                                                       A B
              alleles are codominant because the red blood cells of an I I    that appears in the F 1  generation. With complete domi-
              heterozygote have both kinds of sugars at their surface.  nance, F 1  progeny look like one of the true-breeding par-
              (a)  Codominant lentil coat patterns                 ents. Complete dominance, as we saw in Chapter 2, results
                                                                   in a 3:1 ratio of phenotypes in the F 2 . With incomplete
                                                                   dominance, hybrids resemble neither of the parents and
                                                 D D
                                       S S
                        P             C C       C C                thus  display  neither  pure-breeding  trait.  With  codomi-
                                                                   nance, the phenotypes of both pure-breeding lines show up
                        Gametes       C S        C D               simultaneously in the F 1  hybrid. Both incomplete domi-
                                                                   nance and codominance yield 1:2:1 F 2  ratios.
                                                                   Mendel’s law of segregation still holds
                                       S D
                                                  S D
                        F  (all identical)  C C  C C               The dominance relations of a gene’s alleles do not affect
                         1
                                                                   the alleles’ transmission. Whether two alternative alleles of
                                               C S  C D            a single gene show complete dominance, incomplete domi-
                        F 2                                        nance, or codominance depends on the kinds of proteins
                                           C S                     determined by the alleles and the biochemical function of
                                                S S
                                                      S D
                                               C C   C C           those  proteins  in  the  cell.  These  phenotypic  dominance
                                                                     relations, however, have no bearing on the segregation of
                                           C D                     the alleles during gamete formation.
                                                      D D
                                                S D
                                               C C   C C               As Mendel proposed, cells still carry two copies of
                                   S D
                                                    D D
                      S S
                    1 C C  (spotted) : 2 C C  (spotted/dotted) : 1 C C  (dotted)  each gene, and these copies—a pair of either similar or dis-
                                                                   similar alleles—segregate during gamete formation. Fertil-
              (b)  Codominant blood group alleles
                                                                   ization then restores two alleles to each cell without
              Blood Type                  A                                B                             AB  reference to whether the alleles are the same or different.
                                                                   Variations in dominance relations thus do not detract from
              Red blood
              cell                                        A and B  Mendel’s laws of segregation. Rather, they reflect differ-
                    A sugar         B sugar               sugars   ences in the way gene products control the production of
                                                                   phenotypes, adding a level of complexity to the tasks of
                                                                   interpreting the visible results of gene transmission and
                                                                   inferring genotype from phenotype.
                                   A          B
                                                                   A Gene May Have More Than Two Alleles
                                                 B B
                              A A
                     P                 I I                                         I I
                                                                   Mendel analyzed either-or traits controlled by genes with
                                                                   two alternative alleles, but for many traits, more than two
                                                                   alternatives exist. Here, we look at three such traits: human
                                        A B
                                        I I
                                                                   ABO blood types, lentil seed coat patterns, and   human
                     F 1             AB                              histocompatibility antigens.
                                                                   ABO blood types
              that protrudes from the red blood cell membrane. Each of   If a person with blood type A mates with a person with blood
              the alternative alleles encodes a slightly different form of   type B, it is possible in some cases for the couple to have a
              an enzyme that causes production of a slightly different   child that is neither A nor B nor AB, but a fourth blood type
              form of the complex sugar. In heterozygous individuals, the   called O. The reason? The gene for the ABO blood types has
                                                                                                         A
                                                                                  B
                                                                               A
                                                              B
                                          A
              red blood cells carry both the I -determined and the I -  three alleles: I , I , and i (Fig. 3.5a). Allele I  gives rise to
                                                                                                                   B
              determined sugars on their surface, whereas the cells of   blood type A by specifying an enzyme that adds sugar A, I
                                                            A
              homozygous individuals display the products of either I  or   results in blood type B by specifying an enzyme that adds
               B
              I  alone (Fig. 3.4b). As this example illustrates, when both   sugar B; i does not produce a functional sugar-adding en-
                                                                                      B
              alleles produce a functional gene product, they are usually   zyme. Alleles I  and I  are both dominant to i, and blood
                                                                                A
              codominant for phenotypes analyzed at the molecular level.  type O is therefore a result of homozygosity for allele i.
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