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Metaphase I and Anaphase I
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Metaphase I is marked by the breakdown of the nuclear membrane and the attachment of kinetochore microtubules to the tetrads. Unlike in mitosis, in which sister chromatids are pulled apart because each sister kinetochore is attached to a different pole, both sister kinetochores become attached to spindle microtubules coming from the same pole in metaphase I ( g. 3.23). During anaphase I, cohesin breaks down every place but at the centromeres, allowing sister chromatids to be pulled to
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the same pole: homologous chromosomes are separated ( g. 3.24). This meiotic division is therefore called a reductional division because it reduces the number of chromosomes to half the diploid number in each daughter cell. For every tetrad there is now one chromosome in the form of a chromatid pair, known as a dyad or monovalent, at each pole of the cell. The initial objective of meiosis, separating homologues into different daughter cells, is accomplished. However, since each dyad consists of two sister chromatids, a second, mitosislike division is required to reduce each chromosome to a single chromatid.
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Telophase I and Prophase II
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Depending on the organism, telophase I may or may not be greatly shortened in time. In some organisms, all the expected stages take place; chromosomes enter an interphase con guration as cytokinesis takes place. However, no chromosome duplication (DNA replication) occurs during this abbreviated interphase, termed interkinesis. Next, in these organisms, prophase II begins and
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II. Mendelism and the Chromosomal Theory
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3. Mitosis and Meiosis
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Mitosis and Meiosis
Aster
Kinetochore
Kinetochore microtubule
Figure 3.23 Metaphase of meiosis I; 2n 4. Maternal chromosomes are red; paternal chromosomes are blue. Sister kinetochores (effectively single, merged kinetochores) are attached to microtubules from the same pole.
Anaphase I Figure 3.24 Anaphase of meiosis I; 2n
4. Maternal chromosomes are red; paternal chromosomes are blue. Homologous chromosomes separate and move to opposite poles.
meiosis II proceeds. In still other organisms, the late anaphase I chromosomes go almost directly into metaphase II, virtually skipping telophase I, interphase, and prophase II.
Meiosis II
Meiosis II is basically a mitotic division in which the chromatids of each chromosome are pulled to opposite poles. For each original cell entering meiosis I, four cells emerge at telophase II. Meiosis II is an equational divi-
sion; although it reduces the amount of genetic material per cell by half, it does not further reduce the chromosome number per cell ( g. 3.25). (Sometimes it is simpler to concentrate on the behavior of centromeres during meiosis than on the chromosomes and chromatids. Meiosis I separates maternal from paternal centromeres, and meiosis II separates sister centromeres.) Figure 3.26 summarizes meiosis in corn (Zea mays). In terms of chromosomes, meiosis begins with a diploid cell and produces four haploid cells. In terms of DNA, the process is a bit more complex but has the same
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
3. Mitosis and Meiosis
The McGraw Hill Companies, 2001
Meiosis
Metaphase II
Anaphase II
Telophase II
Interphase Figure 3.25 Meiosis II; 2n
4. Maternal chromosomes are red; paternal chromosomes are blue.
result. Let us call the quantity of DNA in a gamete C. A diploid cell before S phase has 2C DNA, and the same cell after S phase, but before mitosis, has 4C DNA. Mitosis reduces the quantity of DNA to 2C. A cell entering meiosis also has 4C DNA. After the rst meiotic division, each daughter cell has 2C DNA, and after the second meiotic division, each daughter cell has C DNA, the quantity appropriate for a gamete.
The Signi cance of Meiosis
Meiosis is signi cant for several reasons. First, it reduces the diploid number of chromosomes so that each of
four daughter cells has one complete haploid chromosome set. Second, because of the randomness of the process of chromosomal separation, a very large number of different chromosomal combinations can form in the gametes. For example, in human beings, if each gamete could get either the maternal or paternal chromosome, and we have twenty-three chromosomal pairs, 223 or 8,388,608 different combinations can occur.Third, because of crossing over, even more allelic combinations are possible. The process of creating new arrangements, either by crossing over or by independent segregation of homologous pairs of chromosomes, is called recombination. Assuming one hundred thousand genes in a human
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