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2.2 Genetic Analysis According to Mendel   23


                       gamete will carry Y and 1/2 that it will carry y. Because   Figure 2.12  Yellow F 2  peas are of two types: Pure
                       fertilization happens at random, the probability that a par-  breeding and hybrid. The distribution of a pair of contrasting
                       ticular combination of maternal and paternal alleles will   alleles (Y and y) after two generations of self-fertilization. The
                       occur simultaneously in the same zygote is the product of   homozygous individuals of each generation breed true, whereas
                       the independent probabilities of these alleles being pack-  the hybrids do not.
                       aged in egg and sperm. Thus, to find the chance of a Y egg   F 1                Yy
                       (formed as the result of one event) uniting with a Y sperm
                       (the result of an independent event), you simply multiply   Self-
                       1/2 × 1/2 to get 1/4. This is the same fraction of YY progeny   fertilization
                       seen in the Punnett square of Fig. 2.11, which demonstrates   F  YY     Yy              Yy         yy
                       that the Punnett square is simply another way of depicting   2
                       the product rule. It is important to realize that each box
                                                                              Self-
                       in the Punnett square represents an equally likely outcome   fertilization  3:1         3:1
                       of the cross (an equally likely fertilization event)  only
                         because each of the two types of sperm and eggs (Y and y)   F 3  YY  YY  Yy  Yy  yy  YY  Yy  Yy  yy  yy
                       are produced at equal frequencies.                           (All)                                (All)


                       The sum rule                                        kinds of yellow peas (YY and Yy) but only one kind of green
                       While we can describe the moment of random fertilization   pea (yy). In addition, his hypothesis predicted that the YY
                       as the simultaneous occurrence of two independent events,   and Yy yellow peas in the F 2  should be present in a ratio of
                       we can also say that two different fertilization events are   1YY : 2Yy.
                       mutually exclusive. For instance, if Y combines with Y, it   To verify these expectations, Mendel allowed self-
                       cannot also combine with y in the same zygote. A second   fertilization of all the plants in the F 2  generation and
                       rule of probability, the sum rule, states that the probability   counted the types of F 3  progeny (Fig. 2.12). He found
                       of either of two such mutually exclusive events occurring   that the plants that developed from F 2  green peas all pro-
                       is the sum of their individual probabilities. With mutually   duced only green peas in the F 3 , and when the resulting
                       exclusive events:                                   F 3  plants self- fertilized, the next generation (the F 4 ) also
                            Probability of event 1 or event 2 =            produced green peas (not shown). This is what we (and
                                                                           Mendel) would expect of pure-breeding yy lines carrying
                            Probability of event 1 + probability of event 2.  two copies of the recessive allele. The yellow peas were
                          To find the likelihood that an offspring of a Yy hybrid   a different  story. When  Mendel allowed 518  F 2   plants
                       self-fertilization will be a hybrid like the parents, you add   that developed from yellow peas to self-fertilize, he ob-
                       1/4 (the probability of maternal Y uniting with paternal y)   served that 166, roughly 1/3 of the total, were pure-breed-
                       and 1/4 (the probability of the mutually exclusive event   ing yellow through several generations, but the other 352
                       where paternal Y unites with maternal y) to get 1/2, again   (2/3 of the total  yellow F 2  plants) were hybrids because
                       the same result as in the Punnett square.           they gave rise to yellow and green F 3  peas in a ratio of
                          In another use of the sum rule, you could predict the   3:1. Therefore, as Mendel’s theory anticipated, the ratio
                       ratio of yellow to green F 2  progeny. The fraction of F 2  peas   of  YY to  Yy among the 518 F 2  yellow pea plants was
                       that will be yellow is the sum of 1/4 (the event producing     indeed 1:2.
                       YY) plus 1/4 (the mutually exclusive event generating Yy)   It took Mendel years to conduct such rigorous exper-
                       plus 1/4 (the mutually exclusive event producing  yY)   iments on seven pairs of pea traits, but in the end, he was
                       to get 3/4. The remaining 1/4 of the F 2  progeny will be   able to conclude that the segregation of dominant and
                       green. So the yellow-to-green ratio is 3/4 to 1/4, or more   recessive alleles during gamete formation and their ran-
                       simply, 3:1.                                        dom union at fertilization could indeed explain the 3:1
                                                                           ratios he observed whenever he allowed hybrids to
                                                                           self-fertilize. His results, however, raised yet another
                       Further Crosses Verify                              question, one of some importance to future plant and an-
                       the Law of Segregation                              imal breeders. Plants showing a dominant trait, such as
                                                                           yellow peas, can be either pure-breeding (YY) or hybrid
                       The law of segregation was a hypothesis that explained the   (Yy). How can you distinguish one from the other? For
                       data from simple crosses involving monohybrid peas, but   self-fertilizing  plants,  the  answer  is  to  observe  the  ap-
                       Mendel needed to perform additional experiments to check   pearance of the next generation. But how would you dis-
                       its validity. Mendel’s hypothesis, summarized in Fig. 2.11,   tinguish pure-breeding from hybrid individuals in species
                       made the testable prediction that the F 2  should have two   that do not self-fertilize?
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