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PART I  Basic Principles: How Traits Are Transmitted
                                5

                       chapter



                                                   Linkage,



                        Recombination, and


                                   the Mapping of


                                                Genes on

                                                                            Maps illustrate the spatial relationships of objects, such as the
                                    Chromosomes                             locations of subway stations along subway lines. Genetic maps
                                                                            portray the positions of genes along chromosomes.
                                                                            © Rudy Von Briel/PhotoEdit


                                                                            chapter outline

                                                                            •   5.1 Gene Linkage and Recombination
                                                                            •     5.2 Recombination: A Result of Crossing-Over
                                                                              During Meiosis
                       IN  1928,  DOCTORS completed a four-generation pedi-  •     5.3 Mapping: Locating Genes Along a
                       gree tracing two known X-linked traits: red-green color   Chromosome
                       blindness and hemophilia A (the more serious X-linked   •     5.4 The Chi-Square Test and Linkage Analysis
                       form of bleeders disease). The maternal grandfather of   •   5.5 Tetrad Analysis in Fungi
                       the family exhibited both traits, which means that his   •   5.6 Mitotic Recombination and Genetic Mosaics
                       single X chromosome carried mutant alleles of the two
                       corresponding genes. As expected, neither color blind-
                       ness nor hemophilia showed up in his sons and daugh-
                       ters, but two grandsons and one great-grandson inherited both of the X-linked
                       conditions (Fig. 5.1a). The fact that none of the descendants manifested one of
                       the traits without the other suggests that the mutant alleles did not assort inde-
                       pendently during meiosis. Instead they traveled together in the gametes forming
                       one generation and then into the gametes forming the next generation, producing
                       grandsons and great-grandsons with an X chromosome specifying both color
                       blindness and hemophilia. Genes that travel together more often than not exhibit
                       genetic linkage.
                          In contrast, another pedigree following color blindness and the slightly different B
                       form of hemophilia, which also arises from a mutation on the X chromosome, revealed
                       a different inheritance pattern. A grandfather with hemophilia B and color blindness
                       had four grandsons, but only one of them exhibited both conditions. In this family, the
                       genes for color blindness and hemophilia appeared to assort independently, producing
                       in the male progeny all four possible combinations of the two traits—normal vision
                       and normal blood clotting, color blindness and hemophilia, color blindness and normal
                       clotting, and normal vision and hemophilia—in approximately equal frequencies
                       (Fig. 5.1b). Thus, even though the mutant alleles of the two genes were on the same
                       X chromosome in the grandfather, they had to separate to give rise to grandsons III-2
                       and III-3. This separation of genes on the same chromosome is the result of recombi-
                       nation, the occurrence in progeny of new gene combinations not seen in previous
                       generations. (Note that recombinant progeny can result in either of two ways: from the
                       recombination of genes on the same chromosome during gamete formation, discussed

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