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4.5 Gametogenesis   109


                       Figure 4.17  How meiosis contributes to genetic diversity.   essential concepts
                       (a) The variation resulting from the independent assortment of
                       nonhomologous chromosomes increases with the number of   •  In meiosis, chromosomes replicate once (before meiosis I),
                       chromosomes in the genome. (b) Crossing-over between
                       homologous chromosomes ensures that each gamete is unique.  but the nucleus divides twice (meiosis I and II).
                         (a)  Independent assortment      (b)  Recombination  •  During metaphase I, homologous chromosomes connect
                        Orientation I  Orientation II                          to opposite spindle poles. The independent alignment of
                                                                               each pair of homologs ensures the independent
                           A  B           A  b                  C d            assortment of genes carried on different chromosomes.
                                Metaphase I
                           a  b            a  B                 c D          •  Crossing-over during the first meiotic division maintains
                                                              Prophase I       the connection between homologous chromosomes until
                                                   Meiosis I                   anaphase I and contributes to the genetic diversity of
                           A  B            A  b                                gametes.
                                Telophase I                                  •  Sister chromatids separate from each other during
                           a  b            a  B                                meiosis II so that gametes have only one copy of each
                                                                               chromosome.
                                                              Metaphase I    •  Fertilization—the union of egg and sperm—restores the
                                                                               diploid number of chromosomes (2n) to the zygote.
                           A  B            A  b
                                                                             •  Errors during meiosis may produce gametes with
                               Metaphase II                                    missing or extra chromosomes, which often is lethal
                           a  b            a  B                                to offspring.
                                                            C       c  d
                                                  Meiosis II  C  d  c
                            A                                 D      D
                                            A
                             B                b              Metaphase II
                               B               b
                             A               A
                         a                                                   4.5   Gametogenesis
                          b   Telophase II  a  B
                            b              B
                          a              a                                   learning objectives
                                                                             1.  Compare the processes of oogenesis and
                         B
                                        b  A
                          A                   b                      d           spermatogenesis in humans.
                             B  A               A            C     c         2.  Distinguish between the sex chromosome complements
                                                              d
                                         B
                       a                           Gametes                       of human female and male germ-line cells at different
                          b               a
                                b             a              C      D            stages of gametogenesis.
                             a
                                              B                D    c
                                                                           In all sexually reproducing animals, the embryonic germ
                                                                           cells (collectively known as the germ line) undergo a se-
                       Mitosis and Meiosis: A Comparison                   ries of mitotic divisions that yield a collection of special-
                                                                           ized diploid cells, which subsequently divide by meiosis to
                       Mitosis occurs in all types of eukaryotic cells (that is, cells   produce haploid cells. As with other biological processes,
                       with a membrane-bounded nucleus) and is a conservative   many variations on this general pattern have been ob-
                       mechanism that preserves the genetic status quo. Mitosis   served. In some species, the haploid cells resulting from
                       followed by cytokinesis produces growth by increasing the   meiosis are the gametes themselves, while in other spe-
                       number of cells. It also promotes the continual replacement   cies, those cells must undergo a specific plan of differen-
                       of roots, stems, and leaves in plants and the regeneration of   tiation to fulfill that function. Moreover, in certain
                       blood cells, intestinal tissues, and skin in animals.  organisms, the four haploid products of a single meiosis
                          Meiosis, on the other hand, occurs only in sexually   do not all become gametes. Gamete formation, or gameto-
                       reproducing organisms, in just a few specialized germ   genesis, thus gives rise to haploid gametes marked not
                       cells within the reproductive organs that produce haploid   only by the events of meiosis per se but also by cellular
                       gametes. It is not a conservative mechanism; rather, the   events that precede and follow meiosis. Here we illustrate
                       extensive combinatorial changes arising from meiosis are   gametogenesis with a description of egg and sperm forma-
                       one source of the genetic variation that fuels evolution.   tion in humans. The details of gamete formation in several
                       Table 4.3 illustrates the significant contrasts between the   other organisms appear throughout the book in discus-
                       two mechanisms of cell division.                    sions of specific experimental studies.
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