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24     Chapter 2    Mendel’s Principles of Heredity


              Figure 2.13  Genotype versus phenotype in homozygotes   Figure 2.14  How a testcross reveals genotype. An individual
              and heterozygotes. The relationship between genotype and   of unknown genotype, but dominant phenotype, is crossed with a
              phenotype with a pair of contrasting alleles where one allele (Y)   homozygous recessive. If the unknown genotype is homozygous, all
              shows complete dominance over the other (y).         progeny will exhibit the dominant phenotype (cross A). If the unknown
                                                                   genotype is heterozygous, half the progeny will exhibit the dominant
                    Genotype for the Seed    Phenotype             trait, half the recessive trait (cross B).
                    Color Gene
                                                                    Cross A                  Cross B
                            YY
                     Homozygous dominant             Yellow
                                                                    P     YY         yy      P     Yy         yy

                    Dominant    Recessive
                    allele      allele
                            Yy                       Yellow         F 1              y        F 1             y
                        Heterozygous
                                                                                Y   Yy                    Y  Yy
                            yy
                     Homozygous recessive            Green                                                y   yy

                                                                             O spring all yellow  O spring 1:1 yellow to green
              Testcrosses: A way to establish genotype
              Before describing Mendel’s answer, we need to define a   way, the testcross establishes the genotype behind a domi-
              few more terms. An observable characteristic, such as yel-  nant phenotype, resolving any uncertainty.
              low or green pea seeds, is a phenotype, while the actual   As we mentioned earlier, Mendel deliberately simpli-
              pair of alleles present in an individual is its genotype. A YY   fied the problem of heredity, focusing on traits that come in
              or a yy genotype is called homozygous, because the two   only two forms. He was able to replicate his basic monohy-
              copies  of  the  gene  that  determine  the  particular  trait  in   brid findings with corn, beans, and four-o’clocks (plants
              question are the same. In contrast, a genotype with two dif-  with tubular, white or bright red flowers). As it turns out,
              ferent alleles for a trait is heterozygous; in other words, it   his concept of the gene and his law of segregation can be
              is a hybrid for that trait (Fig. 2.13). An individual with a   generalized to almost all sexually reproducing organisms.
              homozygous genotype is a homozygote; one with a hetero-
              zygous genotype is a heterozygote. 
                  Note that the phenotype of a heterozygote (that is, of a   Dihybrid Crosses Reveal the Law
              hybrid) defines which allele is dominant: Because Yy peas   of Independent Assortment
              are yellow, the yellow allele Y is dominant to the y allele for
              green. If you know the genotype and the dominance rela-  Having  determined from monohybrid  crosses  that  genes
              tion of the alleles, you can accurately predict the pheno-  are inherited according to the law of segregation, Mendel
              type. The reverse is not true, however, because some   turned his attention to the simultaneous inheritance of two
              phenotypes can derive from more than one genotype. For   or more apparently unrelated traits in peas. He asked how
              example, the phenotype of yellow peas can result from   two pairs of alleles would segregate in a dihybrid individ-
                either the YY or the Yy genotype.                  ual, that is, in a plant that is heterozygous for two genes at
                  With these distinctions in mind, we can look at the   the same time.
              method Mendel devised for deciphering the unknown gen-   To construct such a dihybrid, Mendel mated true-
              otype. We’ll call it Y–, responsible for a dominant pheno-  breeding plants grown from yellow round peas (YY RR)
              type; the dash represents the unknown second allele, either   with true-breeding plants grown from green wrinkled peas
              Y or y. This method, called the testcross, is a mating in   (yy rr). From this cross he obtained a dihybrid F 1  genera-
              which an individual showing the dominant phenotype, for   tion (Yy Rr) showing the two dominant phenotypes, yellow
              instance, a Y– plant grown from a yellow pea, is crossed   and round (Fig. 2.15). He then allowed these F 1  dihybrids
              with an individual expressing the recessive phenotype, in   to self-fertilize to produce the F 2  generation. Mendel could
              this case a yy plant grown from a green pea. As the Punnett   not predict the outcome of this mating. Would all the F 2
              squares in Fig. 2.14 illustrate, if the dominant phenotype in   progeny be parental types that looked like either the orig-
              question derives from a homozygous YY genotype, all the   inal yellow round parent or the green wrinkled parent? Or
              offspring of the testcross will show the dominant yellow   would some new combinations of phenotypes occur that
              phenotype. But if the dominant parent of unknown geno-  were not seen in the parental lines, such as yellow wrinkled
              type is a heterozygous hybrid (Yy), half of the progeny are   or green round peas? New phenotypic combinations like
              expected to be yellow peas, and the other half green. In this   these are called recombinant types. 
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