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68     Chapter 3    Extensions to Mendel’s Laws


              Figure 3.24  Family pedigrees help unravel the genetic   •  When genes display redundancy for a trait, one dominant
              basis of ocular-cutaneous albinism (OCA). (a) An albino   and normally functioning allele of either gene is sufficient
              Nigerian girl and her sister celebrating the conclusion of the All   to generate the normal phenotype.
              Africa games. (b) A pedigree following the inheritance of OCA in    •  Many traits exhibit locus heterogeneity, in which
              an inbred family indicates that the trait is recessive. (c) A family in
              which two albino parents have nonalbino children demonstrates that   homozygosity for mutations at any one of several genes
              homozygosity for a recessive allele of either of two genes can   can produce the same mutant phenotype.
              cause OCA.                                             •  Complementation occurs in the progeny of pure-breeding
              a: © Radu Sigheti/Reuters                                parents with the same mutant phenotype if the parents
              (a)  Ocular-cutaneous albinism (OCA)                     are homozygous for recessive, nonfunctional alleles of
                                                                       different genes whose products function in a common
                                                                       pathway. 





                                                                    3.3   Extensions to Mendel for
                                                                   Multifactorial Inheritance



                                                                     learning objectives
                                                                     1.  Discuss the factors that can cause different individuals
                                                                        with the same genotype to be phenotypically dissimilar.
              (b)  OCA is recessive                                  2.  Explain how Mendelian genetics is compatible with the
                                                                        fact that many traits, such as human height and skin
               I                                                        colors, exhibit continuous variation.

               II
                                                                   The inheritance of many traits appears to be more complex
              III
                                                                   than can be explained by the participation of only one or two
              IV                                                   genes in patterns compatible with straightforward Mendelian
                                                                   principles. Of course, one reason for this complexity is that
              (c)  Complementation for albinism
                                                                   more than two genes can influence certain traits. But a second
                                                                   reason is that genes are not the only players: The environment
                                                                   and chance events can sometimes exert considerable effects
                      aa BB AA bb                                  on traits that are otherwise genetically determined. In this
                                                         Normal    section we discuss multifactorial traits—traits determined
                                                         Albino    by several different genes, or by the interaction of genes with
                                                                   the environment.

              Aa Bb     Aa Bb       Aa Bb
                                                                   The Same Genotype Does Not Always
                                                                   Produce the Same Phenotype

                                                                   In our discussion of gene interactions so far, we have con-
                                                                   sidered examples in which a genotype reliably fashions a
                essential concepts
                                                                   particular phenotype. But this is not always what happens.
                •  Two genes may interact to affect a single trait; these   Sometimes a genotype is not expressed at all; that is, even
                  interactions may be detected by ratios that can be   though the genotype is present, the expected phenotype
                  predicted from Mendelian principles.             does not appear. Other times, the trait caused by a genotype
                •  Retention of the 9:3:3:1 phenotypic ratio usually indicates   is expressed to varying degrees or in a variety of ways in
                  that two genes function in independent pathways and   different individuals. Factors that alter the phenotypic ex-
                  their alleles interact additively.               pression of genotype include modifier genes, the environ-
                •  In epistasis, an allele at one gene can hide traits   ment, and chance. These factors complicate the interpretation
                  otherwise caused by alleles at another gene.     of breeding experiments.
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