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108    Chapter 4    The Chromosome Theory of Inheritance


              meiotic division. In some species, however, the chromo-    parental cell present at the beginning of this division. For
              somes simply stay condensed. Most importantly, no S   this reason, meiosis II is termed an equational division.
              phase exists during the interphase between meiosis I and
              meiosis II; that is, the chromosomes do not replicate during
              meiotic interphase. The relatively brief interphase between   Mistakes in Meiosis Produce
              meiosis I and meiosis II is known as interkinesis.   Defective Gametes

                                                                   Segregational errors during either meiotic division can
              During Meiosis II, Sister Chromatids                 lead to aberrations, such as trisomies, in the next genera-
              Separate to Produce Haploid Gametes                  tion. If, for example, the homologs of a chromosome pair
                                                                   do not segregate during meiosis I (a mistake known as non-
              The second meiotic division (meiosis II) proceeds in a fash-  disjunction), they may travel together to the same pole and
              ion very similar to that of mitosis, but because the number   eventually become part of the same gamete. Such an error
              of chromosomes in each dividing nucleus has already been   may at fertilization result in any one of a large variety of
              reduced by half, the resulting daughter cells are haploid.   possible trisomies. Most autosomal trisomies in humans,
              The same process occurs in each of the two daughter cells   as we already mentioned, are lethal in utero; one exception
              generated by meiosis I, producing four haploid cells at the   is trisomy 21, the genetic basis of Down syndrome. Like
              end of this second meiotic round (see Fig. 4.15, meiosis II).  trisomy 21, extra sex chromosomes may also be viable but
                                                                   cause a variety of mental and physical abnormalities, such
                                                                   as those seen in Klinefelter syndrome (see Table 4.1).
              Prophase II: The chromosomes condense
              If the chromosomes decondensed during the preceding in-
              terphase, they recondense during prophase II. At the end   Meiosis Contributes to Genetic Diversity
              of prophase II, the nuclear envelope breaks down, and the
              spindle apparatus re-forms.                          The wider the assortment of different gene combinations
                                                                   among members of a species, the greater the chance that at
                                                                   least some individuals will carry combinations of alleles
              Metaphase II: Chromosomes align                      that allow survival in a changing environment. Two aspects
              at the metaphase plate                               of meiosis contribute to genetic diversity in a population.
              The kinetochores of sister chromatids attach to microtubule   First, because only chance governs which paternal or ma-
              fibers emanating from opposite poles of the spindle appa-  ternal homologs migrate to the two poles during the first
              ratus, just as in mitotic metaphase. Nonetheless, two sig-  meiotic division, different gametes carry a different mix of
              nificant features of  metaphase  II distinguish it  from   maternal and paternal chromosomes. Figure 4.17a shows
              mitosis. First, the number of chromosomes is one-half that   how different patterns of homolog migration produce dif-
              in mitotic metaphase of the same species. Second, in most   ferent mixes of parental chromosomes in the gametes. The
              chromosomes, the two sister chromatids are no longer   amount of potential variation generated by this random
              strictly identical because of the recombination through   independent assortment increases with the number of
              crossing-over that occurred during meiosis I. The sister   chromosomes. In  Ascaris, for example, where  n = 2
              chromatids still contain the same genes, but they may carry   (the chromosome complement shown in Fig. 4.17a),
              different combinations of alleles.                   the  random assortment of homologs could produce only
                                                                    2
                                                                   2 , or four types of gametes. In a human being, however,
                                                                                                                23
                                                                   where n = 23, this mechanism alone could generate 2 , or
              Anaphase II: Sister chromatids move                  more than 8 million genetically different kinds of gametes.
              to opposite spindle poles                                A second feature of meiosis, the reshuffling of genetic
              Just as in mitosis, severing of the connection between sister   information through crossing-over during prophase I,
              centromeres allows the sister chromatids to move toward   ensures an even greater amount of genetic diversity in gam-
              opposite spindle poles during anaphase II.           etes. Because crossing-over recombines maternally and pa-
                                                                   ternally derived genes, each chromosome in each different
                                                                   gamete could consist of different combinations of maternal
              Telophase II: Nuclear membranes                      and paternal alleles (Fig. 4.17b).
              re-form, and cytokinesis follows                         Of course, sexual reproduction adds yet another means
              Membranes form around each of four daughter nuclei in   of producing genetic diversity. At fertilization, any one of a
              telophase II, and cytokinesis places each nucleus in a sep-  vast number of genetically diverse sperm can fertilize an egg
              arate cell. The result is four haploid gametes. Note that    with its own distinctive genetic constitution. It is thus not
              at the end of meiosis II, each daughter cell (that is, each   very surprising that, with the exception of identical twins,
              gamete) has the same number of chromosomes as the    the 6 billion people in the world are each genetically unique.
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