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134    Chapter 5    Linkage, Recombination, and the Mapping of Genes on Chromosomes


              (a)  I                                            Figure 5.1  Pedigrees indicate that color blindness and two
                       1       2                                forms of hemophilia are X-linked traits. (a) Transmission of
                 II                                             red-green color blindness and hemophilia A. The traits travel together
                           1         2
                                                                through the pedigree, indicating their genetic linkage. (b) Transmission
                                                                of red-green color blindness and hemophilia B. Even though both genes
                 III
                     1     2    3    4    5    6                are X-linked, the mutant alleles are inherited together in only one of four
                                                                grandsons in generation III. These two pedigrees suggest that the gene
                 IV                                             for color blindness is close to the hemophilia A gene but far away from
                    1        2                    Male          the hemophilia B gene.
                                                  Female
                                                  Hemophilia A  in this chapter, or from the independent assortment of genes on
              (b)  I      1                       Hemophilia B  nonhomologous chromosomes, described in Chapter 4.)
                 II    1      2                                     Two important themes should be kept in mind as we fol-
                                                  Color-blind   low the transmission of genes linked on the same chromo-
                 III                              Hemophilic and  some. The first is that the farther apart two genes are, the
                     1  2   3  4                  color-blind
                                                                greater is the probability of separation through recombination.
                                                                Extrapolating from this general rule, you can see that the gene
                                            for hemophilia A is likely very close to the gene for red-green color blindness because,
                                            as Fig. 5.1a shows, the two rarely separate. By comparison, the gene for hemophilia B
                                            must lie far away from the color blindness gene because, as Fig. 5.1b indicates, new
                                            combinations of alleles of the two genes occur quite often. A second theme is that ge-
                                            neticists can use data about how often genes separate during transmission to map the
                                            genes’ relative locations on a chromosome. Such mapping is a key to sorting out and
                                            tracking down the components of complex genetic networks; it is also crucial to ge-
                                            neticists’ ability to isolate and  characterize genes at the molecular level.





               5.1   Gene Linkage                                  Some Genes on the Same Chromosome
              and Recombination                                    Do Not Assort Independently—Instead,
                                                                   They Are Linked

                                                                   We begin our analysis with X-linked Drosophila genes be-
                learning objectives                                cause they were the first to be assigned to a specific chro-
                                                                   mosome.  As  we  outline  various  crosses,  remember  that
                1.  Define linkage with respect to gene loci and
                   chromosomes.                                    females carry two X chromosomes, and thus two alleles for
                2.  Differentiate between parental and recombinant   each X-linked gene. Males, in contrast, have only a single
                   gametes.                                        X chromosome (from the female parent), and thus only a
                3.  Conclude from ratios of progeny in a dihybrid cross   single allele for each of these genes.
                                                                       We look first at two X-linked genes that determine a
                   whether two genes are linked.                   fruit fly’s eye color and body color. These two genes are
                4.  Explain how a testcross can provide evidence for or   said to be syntenic because they are located on the same
                   against linkage.
                                                                   chromosome. The white gene was introduced in Chapter 4;
                                                                                                              +
                                                                   you will recall that the dominant wild-type allele w  speci-
                                                                   fies red eyes, while the recessive mutant allele w confers
                                                                                                                   +
              If people have roughly 27,000 genes but only 23 pairs of   white eyes. The alleles of the yellow body color gene are y
              chromosomes, most human chromosomes must carry       (the dominant wild-type allele for brown bodies) and y (the
                hundreds, if not thousands, of genes. This is certainly true   recessive mutant allele for yellow bodies). To avoid confu-
                                                                                                +
              of the human X chromosome, which contains about 1100   sion, note that lowercase y and y  refer to alleles of the
              protein-coding genes, as just described in Chapter 4. Rec-  yellow gene, while capital Y refers to the Y chromosome
              ognition that many genes reside on each chromosome   (which does not carry genes for either eye or body color).
              raises an important question. If genes on different chromo-  You should also pay attention to the slash symbol (/), which
              somes assort independently because nonhomologous chro-  is used to separate genes found on the two chromosomes of
              mosomes align independently on the spindle during meiosis   a pair (either the X and Y chromosomes as in this case, or
              I, how do genes on the same chromosome assort?       a pair of X chromosomes or homologous autosomes). Thus
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