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2.2 Genetic Analysis According to Mendel   25


                       Figure 2.15  A dihybrid cross produces parental types   for pea color and for pea shape assort independently, the
                       and recombinant types. In this dihybrid cross, pure-breeding   allele for pea shape in a gamete carrying Y could with equal
                       parents (P) produce a genetically uniform generation of F 1  dihybrids.   likelihood be either R or r. Thus, the presence of a particu-
                       Self-pollination or cross-pollination of the F 1  plants yields the   lar allele of one gene, say, the dominant Y for pea color,
                       characteristic F 2  phenotypic ratio of 9:3:3:1.
                                                                           provides no information whatsoever about the allele of the
                       P                                                   second gene. Each dihybrid of the F 1  generation can there-
                                     YY RR           yy rr                 fore make four kinds of gametes: Y R, Y r, y R, and y r. In a
                                                                           large number of gametes, the four kinds will appear in an
                                                                           almost perfect ratio of 1:1:1:1, or put another way, roughly
                       Gametes        Y R            y r                   1/4 of the eggs and 1/4 of the sperm will contain each of the
                                                                           four possible combinations of alleles. That “the different
                                                                           kinds of germinal cells [eggs or sperm] of a hybrid are pro-
                       F  (all identical)                                  duced on the average in equal numbers” was yet another
                        1
                                     Yy Rr          Yy Rr                  one of Mendel’s incisive insights.
                                                                               At fertilization then, in a mating of dihybrids, 4 differ-
                       F 2                         1/4  1/4  1/4   1/4     ent kinds of eggs can each combine with any 1 of 4 different
                                                                           kinds of sperm, producing a total of 16 possible zygotes.
                                                  Y R   Y r  y R   y r
                                                                           Once again, a Punnett square is a convenient way to visual-
                                                                           ize the process (Fig. 2.15). Using the same kind of logic
                                          1/4  Y R
                                                 YY RR YY Rr Yy RR Yy Rr   previously applied to the Punnett square for monohybrid
                                                                           crosses (review Fig. 2.11), each of the 16 boxes with colored
                                          1/4  Y r                         peas in the Punnett square for the dihybrid cross in Fig. 2.15
                                                  YY Rr  YY rr   Yy Rr  Yy rr
                                                                           represents an equally likely fertilization event. Again, each
                                          1/4  y R                         box is an equally likely outcome only because each of the
                                                 Yy RR  Yy Rr  yy RR  yy Rr  different gamete types is produced at equal frequency in

                                          1/4  y r                         each parent. Therefore, using the product rule, the frequency
                                                  Yy Rr  Yy rr  yy Rr  yy rr  of the progeny type in each box is 1/4 × 1/4 = 1/16.
                                                              Each box:        If you look at the square in Fig. 2.15, you will see that
                                                              1/4 × 1/4 = 1/16  some of the 16 potential allelic combinations are identical.
                         Type    Genotype    Phenotype  Number Phenotypic   In fact, only nine different genotypes exist—YY RR, YY Rr,
                                                                 Ratio
                                                                           Yy RR, Yy Rr, yy RR, yy Rr, YY rr, Yy rr, and yy rr—because
                       Parental   Y– R–           yellow round  315  9/16  the source of the alleles (egg or sperm) does not make any
                                                                           difference. If you look at the combinations of traits
                       Recombinant  yy R–           green round  108  3/16    determined by the nine genotypes, you will see only four
                                                                           phenotypes—yellow round, green round, yellow wrinkled,
                                                                           and green wrinkled—in a ratio of 9:3:3:1. If, however, you
                       Recombinant  Y– rr           yellow wrinkled  101  3/16
                                                                           look only at pea color or only at pea shape, you can see that
                                                                           each trait is inherited in the 3:1 ratio predicted by Mendel’s
                       Parental   yy rr           green wrinkled  32  1/16
                                                                           law of segregation. In the Punnett square, there are 12 yel-
                                                                           low for every 4 green and 12 round for every 4 wrinkled. In
                       Ratio of yellow (dominant) to green (recessive)   =    12:4 or 3:1
                                                                           other words, the ratio of each dominant trait (yellow or
                       Ratio of round (dominant) to wrinkled (recessive)  =    12:4 or 3:1
                                                                           round) to its antagonistic recessive trait (green or wrinkled)
                                                                           is 12:4, or 3:1. This means that the inheritance of the gene
                          When Mendel counted the F 2  generation of one exper-  for pea color is unaffected by the inheritance of the gene for
                       iment, he found 315 yellow round peas, 108 green round,   pea shape, and vice versa.
                       101 yellow wrinkled, and 32 green wrinkled. Both yellow   The preceding analysis became the basis of Mendel’s
                       wrinkled and green round recombinant phenotypes did, in   second general genetic principle, the law of independent
                       fact, appear, providing evidence that some shuffling of the     assortment:  During gamete formation, different pairs of
                       alleles of different genes had taken place.           alleles segregate independently of each other (Fig. 2.16). The
                                                                           independence of their segregation and the subsequent  random
                                                                           union of gametes at fertilization determine the phenotypes
                       The law of independent assortment                   observed. Using the product rule for assessing the probability
                       From the observed ratios, Mendel inferred the biological   of independent events, you can see mathematically how the
                       mechanism of that shuffling—the independent assortment   9:3:3:1 phenotypic ratio observed in a dihybrid cross derives
                       of gene pairs during gamete formation. Because the genes   from two separate 3:1 phenotypic ratios. If the two sets of
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