Page 23 - Applied Probability
P. 23

1. Basic Principles of Population Genetics
                              6
                                           1
                                                    1



                                                             2
                                        2
                                                     2p 1 p 2 + p
                                                                 =2p 1(p 1 + p 2 )p 2 (p 1 + p 2 )
                                    2 p + 2p 1 p 2
                                        1
                                                             2
                                                    2
                                           2
                                                                 =2p 1p 2
                                                   1
                                                              2


                                                            2

                                                                      p 2 (p 1 + p 2 )
                                                                 =
                                                    2p 1 p 2 + p
                                                            2
                                                   2
                                                                      2
                                                                 = p .           2
                                                                      2
                              Thus, after a single round of random mating, genotype frequencies stabilize
                              at the Hardy-Weinberg proportions.
                                We may deduce the same result by considering the gamete population.
                              A 1 gametes have frequency p 1 and A 2 gametes frequency p 2 . Since random
                              union of gametes is equivalent to random mating, A 1 /A 1 is present in the
                                                           2
                              next generation with frequency p , A 1 /A 2 with frequency 2p 1p 2 , and A 2 /A 2
                                                           1
                                             2
                              with frequency p . In the gamete pool from this new generation, A 1 again
                                             2
                                                   2
                              occurs with frequency p + p 1 p 2 = p 1 (p 1 + p 2 )= p 1 and A 2 with frequency
                                                   1
                              p 2 . In other words, stability is attained in a single generation. This random
                              union of gametes argument generalizes easily to more than two alleles.
                                Hardy-Weinberg equilibrium is a bit more subtle for X-linked loci. Con-
                              sider a locus on the X chromosome and any allele at that locus. At genera-
                              tion n let the frequency of the given allele in females be q n and in males be
                              r n . Under our stated assumptions for Hardy-Weinberg equilibrium, one can
                                                                              2
                                                                                   1
                              show that q n and r n converge quickly to the value p = q 0 + r 0 . Twice as
                                                                              3    3
                              much weight is attached to the initial female frequency since females have
                              two X chromosomes while males have only one.
                                Because a male always gets his X chromosome from his mother, and his
                              mother precedes him by one generation,
                                                             = q n−1 .                     (1.1)
                                                         r n
                              Likewise, the frequency in females is the average frequency for the two sexes
                              from the preceding generation; in symbols,
                                                            1       1
                                                        =    q n−1 + r n−1 .               (1.2)
                                                    q n
                                                            2       2
                              Equations (1.1) and (1.2) together imply
                                           2     1        2 1       1         1

                                            q n + r n  =      q n−1 + r n−1 + q n−1
                                           3     3        3 2       2         3
                                                          2       1
                                                      =    q n−1 + r n−1 .                 (1.3)
                                                          3       3
                                                               2
                                                                    1
                              It follows that the weighted average q n + r n = p for all n.
                                                               3    3
                                From equations (1.2) and (1.3), we deduce that
                                                     3    1
                                      q n − p = q n − p + p
                                                     2    2
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