During interphase (IN-ter-fa¯ z) the cell replicates its DNA
through a process that will be described shortly. It also produces additional organelles and cytosolic components in anticipation of cell division. Interphase is a state of high metabolic activity; it is during this time that the cell does most of its growing. Interphase consists of three phases: G1, S, and G2 (Figure 3.30). The S stands for synthesis of DNA. Because the G phases are periods when there is no activity related to DNA duplication, they are thought of as gaps or interruptions in DNA duplication. The G1 phase is the interval between the mitotic phase and the S phase. During G1, the cell is metabolically active; it replicates most of its organelles and cytosolic components but not its DNA. Replication of centrosomes also begins in the G1 phase. Virtually all the cellular activities described in this chapter happen during G1. For a cell with a total cell cycle time of 24 hours, G1 lasts 8 to 10 hours. However, the duration of this phase is quite variable. It is very short in many embryonic cells or cancer cells. Cells that remain in G1 for a very long time, perhaps destined never to divide again, are said to be in the G0 phase. Most nerve cells are in the G0 phase. Once a cell enters the S phase, however, it is committed to go through the rest of the cell cycle. The S phase, the interval between G1 and G2, lasts about 8 hours. During the S phase, DNA replication occurs. As a result of DNA replication, the two identical cells formed during cell division later in the cell cycle will have the same genetic material. The G2 phase is the interval between the S phase and the mitotic phase. It lasts 4 to 6 hours. During G2, cell growth continues, enzymes and other proteins are synthesized in preparation for cell division, and replication of centrosomes is completed. When DNA replicates during the S phase, its helical structure partially uncoils, and the two strands separate at the points where hydrogen bonds connect base pairs (Figure 3.31). Each exposed base of the old DNA strand then pairs with the complementary base of a newly synthesized nucleotide. A new DNA strand takes shape as chemical bonds form between neighboring nucleotides. The uncoiling and complementary base pairing continues until each of the two original DNA strands is joined with a newly formed complementary DNA strand. The original DNA molecule has become two identical DNA molecules. Mitotic Phase The mitotic (M) phase of the cell cycle, which results in the formation of two identical cells, consists of a nuclear division (mitosis) and a cytoplasmic division (cytokinesis) to form two identical cells. The events that occur during mitosis and cytokinesis are plainly visible under a microscope because chromatin condenses into discrete chromosomes. NUCLEAR DIVISION: MITOSIS Mitosis, as noted earlier, is the distribution of two sets of chromosomes into two separate nuclei. The process results in the exact partitioning of genetic information. For convenience, biologists divide the process into four stages: prophase, metaphase, anaphase, and telophase. However, mitosis is a continuous process; one stage merges seamlessly into the next. 1. Prophase (PRO¯ -fa¯ z). During early prophase, the chromatin fibers condense and shorten into chromosomes that are visible under the light microscope (Figure 3.32b). The condensation process may prevent entangling of the long DNA strands as they move during mitosis. Because longitudinal DNA replication took place during the S phase of interphase, each prophase chromosome consists of a pair of identical strands called chromatids. A constricted region called a centromere (SEN-tro¯- me¯ r) holds the chromatid pair together. At the outside of each centromere is a protein complex known as the kinetochore (ki-NET-o¯-kor). Later in prophase, tubulins in the pericentriolar material of the centrosomes start to form the mitotic spindle, a football-shaped assembly of microtubules that attach to the kinetochore (Figure 3.32b). As the microtubules lengthen, they push the centrosomes to the poles (ends) of the cell so that the spindle extends from pole to pole. The mitotic spindle is responsible for the separation of chromatids to opposite poles of the cell. Then, the nucleolus disappears and the nuclear envelope breaks down. 2. Metaphase (MET-a-fa¯z). During metaphase, the microtubules of the mitotic spindle align the centromeres of the chromatid pairs at the exact center of the mitotic spindle (Figure 3.32c). This midpoint region is called the metaphase plate. 3. Anaphase (AN-a-fa¯ z). During anaphase, the centromeres split, separating the two members of each chromatid pair, which move toward opposite poles of the cell (Figure 3.32d). Once separated, the chromatids are termed chromosomes. As the chromosomes are pulled by the microtubules of the mitotic spindle during anaphase, they appear V-shaped because the centromeres lead the way, dragging the trailing arms of the chromosomes toward the pole. 4. Telophase (TEL-o¯-fa¯z). The final stage of mitosis, telophase, begins after chromosomal movement stops (Figure 3.32e). The identical sets of chromosomes, now at opposite poles of the cell, uncoil and revert to the threadlike chromatin form. A nuclear envelope forms around each chromatin mass, nucleoli reappear in the identical nuclei, and the mitotic spindle breaks up.