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26     Chapter 2    Mendel’s Principles of Heredity


              Figure 2.16  The law of independent assortment. In a     sequence of columns. In Fig. 2.17, the first column shows
              dihybrid cross, each pair of alleles assorts independently during   the two possible pea color phenotypes; the second column
              gamete formation. In the gametes, Y is equally likely to be found with   demonstrates that each pea color can occur with either of
              R or r (that is, Y R = Y r); the same is true for y (that is, y R = y r). As   two pea shapes. Again, the 9:3:3:1 ratio of phenotypes is
              a result, all four possible types of gametes (Y R, Y r, y R, and y r) are
              produced in equal frequency among a large population.  apparent.  You will see later that branched-line diagrams
                                                                   are more convenient than Punnett squares for predicting the
                                            Possible allele
                        Alleles in   Gamete                        outcomes of crosses involving more than two genes.
                        parental cell  formation  combinations
                                            in gametes
                                                                   Testcrosses with dihybrids
                                               Y R 1/4
                                                                   An understanding of dihybrid crosses has many applica-
                                                                   tions. Suppose, for example, that you work for a nursery
                                               Y r 1/4             that has three pure-breeding strains: yellow wrinkled, green
                           Y
                           y                                       round, and green  wrinkled. Your assignment is to grow
                           R                                       pure-breeding plants guaranteed to produce yellow round
                           r                   y R 1/4
                                                                   peas. How would you proceed? 
                                                                       One answer is to cross your two pure-breeding strains
                                               y r 1/4             (YY rr  ×  yy RR) to generate a dihybrid (Yy Rr). Then self-
                                                                   cross the dihybrid and plant only the yellow round peas.
                                                                   Only one out of nine of such progeny—those grown from
              alleles assort independently, the yellow-to-green ratio in the   peas with a YY RR genotype—will be appropriate for your
              F 2  generation will be 3/4 : 1/4, and likewise, the round-to-  uses. To find these plants, you could subject each yellow
              wrinkled ratio will be 3/4 : 1/4. To find the probability that   round candidate to a testcross for genotype with a green
              two independent events such as yellow and round will occur   wrinkled (yy rr) plant, as illustrated in  Fig. 2.18. If the
              simultaneously in the same plant, you multiply as follows:
                  Probability of yellow round = 3/4 × 3/4 = 9/16   Figure 2.18  Testcrosses with dihybrids. Testcrosses involving
                                                                   two pairs of independently assorting alleles yield different, predictable
                  Probability of green round = 1/4 × 3/4 = 3/16    results depending on the tested individual’s genotype for the two
                                                                   genes in question.
                  Probability of yellow wrinkled = 3/4 × 1/4 = 3/16
                                                                   Cross A                     Cross B
                  Probability of green wrinkled = 1/4 × 1/4 = 1/16
                                                                     P   YY RR       yy rr       P  YY Rr       yy rr
                  Thus, in a population of F 2  plants, there will be a
              9:3:3:1 phenotypic ratio of yellow round to green round to
              yellow wrinkled to green wrinkled.                     F 1              y r       F 1             y r


              Branched-line diagrams                                            Y R                        Y R
                                                                                    Yy Rr                      Yy Rr
              A convenient way to keep track of the probabilities of each
              potential outcome in a genetic cross is to construct a                                       Y r
              branched-line diagram (Fig. 2.17), which shows all the                                            Yy rr
              possible genotypes or phenotypes for each gene in a   Cross C                    Cross D
              Figure 2.17  Following crosses with branched-line       P   Yy RR      yy rr       P  Yy Rr       yy rr
              diagrams. A branched-line diagram, which uses a series of
              columns to track every gene in a cross, provides an organized   F 1                F 1
              overview of all possible outcomes. This branched-line diagram of a      y r                       y r
              dihybrid cross generates the same phenotypic ratios as the Punnett
              square in Fig. 2.15, showing that the two methods are equivalent.
                                                                                Y R                        Y R
               Gene 1           Gene 2          Phenotypes                          Yy Rr                      Yy Rr
                                                                                y R                        Y r
                                3/4 round       9/16 yellow round
               3/4 yellow                                                            yy Rr                     Yy rr
                                1/4 wrinkled    3/16 yellow wrinkled
                                                                                                           y R
                                                                                                                yy Rr
                                3/4 round       3/16 green round
               1/4 green                                                                                   y r
                                1/4 wrinkled    1/16 green wrinkled                                             yy rr
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