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12. Models of Recombination
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                              12.7 Application to Drosophila Data
                              As an application of the various recombination models, we now briefly dis-
                              cuss the classic Drosophila data of Morgan et al. [20]. These early geneticists
                              phenotyped 16,136 flies at 9 loci covering almost the entire Drosophila X
                              chromosome. Because of the nature of the genetic cross employed, a fly
                              corresponds to a gamete scorable on each interlocus interval as recombi-
                              nant or nonrecombinant. Table 12.1 presents the gamete counts n recorded.
                              Here the set S denotes the recombinant intervals. For example, 6,607 flies
                              were nonrecombinant on all intervals (the first category), and one fly was
                              recombinant on intervals 5, 6, and 8 and nonrecombinant on all remaining
                              intervals (the last category).
                                        TABLE 12.1. Gamete Counts in the Morgan et al. Data
                                       S      n      S      n      S      n     S      n
                                       φ     6607   {2, 4}  38   {6, 7}  21   {2, 5, 6}  3
                                      {1}     506   {2, 5}  85   {6, 8}  30   {2, 5, 7}  4
                                      {2}    1049   {2, 6}  237  {7, 8}   2   {2, 5, 8}  1
                                      {3}     855   {2, 7}  123  {1, 2, 3}  1  {2, 6, 7}  2
                                      {4}    1499   {2, 8}  70  {1, 2, 6}  1  {2, 6, 8}  3
                                      {5}     937   {3, 4}  22  {1, 3, 5}  1  {2, 7, 8}  2
                                      {6}    1647   {3, 5}  55  {1, 4, 5}  1  {3, 4, 7}  2
                                      {7}     683   {3, 6}  177  {1, 4, 6}  1  {3, 4, 8}  1
                                      {8}     379   {3, 7}  88  {1, 4, 7}  2  {3, 5, 6}  1
                                      {1, 2}    3   {3, 8}  38  {1, 4, 8}  1  {3, 5, 7}  2
                                      {1, 3}    6   {4, 5}  41  {1, 5, 7}  2  {3, 5, 8}  3
                                      {1, 4}   41   {4, 6}  198  {1, 5, 8}  1  {3, 6, 7}  1
                                      {1, 5}   55   {4, 7}  159  {1, 6, 8}  1  {3, 6, 8}  1
                                      {1, 6}  118   {4, 8}  91  {2, 3, 6}  1  {4, 5, 8}  1
                                      {1, 7}   54   {5, 6}  35  {2, 4, 6}  4  {4, 6, 8}  4
                                      {1, 8}   34   {5, 7}  49  {2, 4, 7}  5  {4, 7, 8}  1
                                      {2, 3}    3   {5, 8}  40  {2, 4, 8}  6  {5, 6, 8}  1


                                Table 12.2 summarizes the results presented in the papers [18, 23]. Hal-
                              dane’s model referred to in the first row of the table fits the data poorly.
                              The count-location model yields an enormous improvement in the maxi-
                              mum loglikelihood displayed in the last column of the table. In the count-
                              location model, the maximum likelihood estimates of the count proba-
                              bilities are (q 0 ,q 1 ,q 2 ,q 3 )=(.06,.41,.48,.05); these estimates correct the
                              slightly erroneous values given in [23]. The departure of the count proba-
                              bilities from a Poisson distribution is one of the reasons Haldane’s model
                              fails so miserably. The chi-square and mixture models referred to in Table
                              12.2 are special cases of the Poisson-skip model. The best fitting chi-square
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