第11章減数分裂と有性生殖

During the meiotic interphase, each chromosome is duplicated. The sister chromatids that are formed during synthesis are held together at the centromere region by cohesin proteins. All chromosomes are attached to the nuclear envelope by their tips. As the cell enters prophase I, the nuclear envelope begins to fragment and the proteins holding homologous chromosomes locate each other. The four sister chromatids align lengthwise, and a protein lattice called the synaptonemal complex is formed between them to bind them together. The synaptonemal complex facilitates crossover between nonsister chromatids, which is observed as chiasmata along the length of the chromosome. As prophase I progresses, the synaptonemal complex breaks down and the sister chromatids become free, except where they are attached by chiasmata. At this stage, the four chromatids are visible in each homologous pairing and are called a tetrad.

Random alignment leads to new combinations of traits. The chromosomes that were originally inherited by the gamete-producing individual came equally from the egg and the sperm. In metaphase I, the duplicated copies of these maternal and paternal homologous chromosomes line up across the center of the cell. The orientation of each tetrad is random. There is an equal chance that the maternally derived chromosomes will be facing either pole. The same is true of the paternally derived chromosomes. The alignment should occur differently in almost every meiosis. As the homologous chromosomes are pulled apart in anaphase I, any combination of maternal and paternal chromosomes will move toward each pole. The gametes formed from these two groups of chromosomes will have a mixture of traits from the individual’s parents. Each gamete is unique.

In metaphase I, the homologous chromosomes line up at the metaphase plate. In anaphase I, the homologous chromosomes are pulled apart and move to opposite poles. Sister chromatids are not separated until meiosis II. The fused kinetochore formed during meiosis I ensures that each spindle microtubule that binds to the tetrad will attach to both sister chromatids.

All of the stages of meiosis I, except possibly telophase I, are unique because homologous chromosomes are separated, not sister chromatids. In some species, the chromosomes do not decondense and the nuclear envelopes do not form in telophase I. All of the stages of meiosis II have the same events as the stages of mitosis, with the possible exception of prophase II. In some species, the chromosomes are still condensed and there is no nuclear envelope. Other than this, all processes are the same.

The chromosomes of the individual cannot cross over during meiosis if the individual cannot make recombination nodules. This limits the genetic diversity of the individual’s gametes to what occurs during independent assortment, with all daughter cells receiving complete maternal or paternal chromatids. An individual who cannot produce diverse offspring is considered less fit than individuals who do produce diverse offspring.

Crossing over does not occur during prophase II; it only occurs during prophase I. In prophase II, there are still two copies of each gene, but they are on sister chromatids within a single chromosome (rather than homologous chromosomes as in prophase I). Therefore, any crossover event would still produce two identical chromatids. Because it is advantageous to avoid wasting energy on events that will not increase genetic diversity, crossing over does not occur.