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II. Mendelism and the Chromosomal Theory
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3. Mitosis and Meiosis
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The McGraw Hill Companies, 2001
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Mitosis and Meiosis
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Table 3.3 Chromosome Number for Selected
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Species
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Species Human being (Homo sapiens) Garden pea (Pisum sativum) Fruit y (Drosophila melanogaster) House mouse (Mus musculus) Roundworm (Ascaris sp.) Pigeon (Columba livia) Boa constrictor (Constrictor constrictor) Cricket (Gryllus domesticus) Lily (Lilium longi orum) Indian fern (Ophioglossum reticulatum) 2n 46 14 8 40 2 80 36 22 24 1,260
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Note: 2n is the diploid complement. The fern has the highest known diploid chromosome number.
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is less organized. In animals, a constriction of the cell membrane distributes the cytoplasm. In plants, the growth of a cell plate accomplishes the same purpose.
THE CELL CYCLE
The continuity of life depends on cells growing, replicating their genetic material, and then dividing, a process called the cell cycle ( g. 3.6). Although cells usually divide when they have doubled in volume, the control of this process is very complex and precise. Not only do all the steps have to occur in sequence, but the cell must also know when to proceed and when to wait. Continuing at inappropriate moments for example, before the DNA has replicated or when the chromosomes or spindle are damaged could have catastrophic consequences to a cell or a whole organism. Numerous stops occur during the cycle to assess whether the next step should proceed. Early research into the cell cycle involved fusing cells in different stages of the cycle (such as the G1, S, and G2 phases; see g. 3.6) to determine whether the cytoplasmic components of one cell would affect the behavior of the other. Results of these experiments led to the discovery of a protein complex called the maturationpromoting factor (MPF) because of its role in causing oocytes to mature. It is now also referred to as the mitosis-promoting factor since it initiates the mitosis phase of the cell cycle. Further research has shown that MPF is made of two proteins, one that oscillates in quantity during the cell cycle and one whose quantity is con-
stant. The oscillating component is referred to as cyclin; the constant gene product is an enzyme controlled by the cdc2 gene (cdc stands for cell division cycle) called Cdc2p. Cdc2p is a kinase, an enzyme that phosphorylates other proteins, transferring a phosphate group from ATP to an amino acid of the protein it is acting on. (Phosphorylation controls many of the processes in mitosis and in metabolism in general; for example, the nuclear membrane begins to break down when its subunits are phosphorylated.) Because the Cdc2p kinase works when combined with cyclin, it is referred to as a cyclindependent kinase (CDK). Several of these kinasecyclin combinations control stages of the cell cycle; the cyclin of the mitosis-promoting factor is called cyclin B. In general, cylin-dependent kinases are regulated by phosphorylation and dephosphorylation, cyclin levels, and activation or deactivation of inhibitors. Normally, Cdc2p remains at high levels in the cell but does not initiate mitosis for two reasons. First, phosphate groups block its active site, the place on the enzyme that actually does the phosphorylating. Second, the enzyme can only function when it combines with a molecule of cyclin B, the protein that oscillates during the cell cycle. Cyclin B is at very low levels when mitosis ends. During ensuing cell growth, numbers of cyclin B molecules increase, combining with Cdc2p proteins until a critical quantity is reached. However, Cdc2p-cyclin B complexes are still not active. That requires the product of another gene to dephosphorylate the Cdc2p-cyclin B complex. At that point, the Cdc2p-cyclin B complex goes into action, initiating the changes that begin mitosis ( g. 3.7). Presumably the cell is ready for mitosis at that point, having
Gap (G2) DNA synthesis (S) 40% Mitosis (M) 10% Gap (G1) 25% 25%
Figure 3.6 Cell cycle in the broad bean, Vicia faba. Total time in the cycle is under twenty hours. The DNA content of the cell doubles during the S phase and is then reduced back to its original value by mitosis.
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