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5.3 Mapping: Locating Genes Along a Chromosome   151


                       analyses reveal that most chromosomes in human sperm   been assigned to a particular chromosome, the terms chro-
                       have undergone only a single crossover.             mosome and linkage group become synonymous. If you can
                          Drosophila provides an extreme example: No recom-  demonstrate that gene A is linked to gene B, B to C, C to D,
                       bination occurs during meiosis in males. If you review the   and D to E, you can conclude that all these genes are syn-
                       examples already discussed in this chapter, you will dis-  tenic. When the genetic map of a genome becomes so dense
                       cover that they all measure recombination among the   that it is possible to show that any gene on a chromosome is
                         progeny of doubly heterozygous Drosophila females. Prob-  linked to another gene on the same chromosome, the number
                       lem 19 at the end of this chapter shows how geneticists can   of linkage groups equals the number of pairs of homologous
                       exploit the absence of recombination in Drosophila males   chromosomes in the species. Humans have 23 linkage
                       to establish rapidly that genes far apart on the same   groups, mice have 20, and fruit flies have 4 (Fig. 5.18).
                         chromosome are indeed syntenic.                       The total genetic distance along a chromosome, which
                                                                           is obtained by adding many short distances between genes,
                       Multiple-Factor Crosses Help Establish              may be much more than 50 m.u. For example, the two long
                       Linkage Groups                                      Drosophila autosomes are both slightly more than 100 m.u.
                                                                           in length (Fig. 5.18), while the longest human chromosome
                       Genes chained together by linkage relationships are known   is approximately 270 m.u. Recall, however, that even with
                       collectively as a linkage group. When enough genes have   the longest chromosomes, pairwise crosses between genes



                       Figure 5.18  Drosophila melanogaster has four linkage groups. A genetic map of the fruit fly, showing the position of many
                       genes affecting body morphology, including those used as examples in this chapter (highlighted in bold). Because so many Drosophila genes
                       have been mapped, each of the four chromosomes can be represented as a single linkage group.

                                           X chromosome

                                                                                               bent wing
                                                                net veins                      cubitus veins
                                               yellow body  0.0  artistaless antenna  0.0  roughoid eyes 0.0
                                          0.0  scute bristles  1.3  star eyes  0.2  veinlet veins  shaven hairs
                                                                                               grooveless scutellum
                                          1.5  white eyes                                      eyeless
                                          3.0  facet eyes  4.0  held-out wings
                                          5.5  echinus eyes
                                          7.5  ruby eyes
                                          13.7            13.0  dumpy wings
                                               crossveinless wings
                                                          16.5  thick veins
                                         20.0  cut wings                   19.2  javelin bristles
                                          21.0  singed bristles
                                                                           26.0  sepia eyes
                                          27.7  lozenge eyes               26.5  hairy body
                                         33.0  vermilion eyes
                                          36.1  miniature wings
                                                                           41.0  dichaete bristles
                                         43.0  sable body                  43.2  thread arista
                                         44.0  garnet eyes  48.5  black body  44.0    scarlet eyes
                                                          51.0  reduced bristles  48.0  pink eyes
                                         55.0  rudimentary  54.5  purple eyes  52.0  rosy eyes
                                         56.7  forked bristles  54.8  short bristles  58.2  stubble bristles
                                         57.0  bar eyes   55.0  light eyes  58.5  spineless bristles
                                         59.5  fused veins  57.5  cinnabar eyes  58.7  bithorax body
                                         62.5  carnation eyes              62.0  stripe body
                                         66.0  bobbed hairs  66.7  scabrous eyes  63.0  glass eyes
                                                          67.0  vestigial wings  66.2  delta veins
                                                          72.0  lobe eyes  69.5  hairless bristles
                                                                           70.7  ebony body
                                                          75.5  curved wings
                                                                           74.7  cardinal eyes


                                                                            91.1  rough eyes


                                                          100.5  plexus wings  100.7  claret eyes
                                              Wild type
                                                          104.5  brown eyes
                                                          107.0  blistered wings  106.2  minute bristles
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