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Tamarin: Principles of Genetics, Seventh Edition
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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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The Cell Cycle
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gone through G1, S, and G2 phases (which we will discuss in detail later in the chapter). Once mitosis has been initiated, cyclin B, along with other proteins that have served their purpose by this point in the cell cycle, breaks down with the help of a protein complex called the anaphase-promoting complex (APC), also called the cyclosome. The cyclosome works by attaching a ubiquitin molecule to the proteins that are to be broken down. (Ubiquitin is a polypeptide of 76 amino acids; it directs the attached protein into a breakdown pathway discussed in chapter 16.) Cdc2p is then phosphorylated to block its active site.The cell now completes mitosis and enters G1; quantities of cyclin B are very low, and virtually no functioning Cdc2p-cyclin B remains ( g. 3.7). Thus, active Cdc2p is the kinase that controls the initiation of mitosis. Some points in the cell cycle, such as the initiation of mitosis, can be delayed until all necessary conditions are
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in place. These checkpoints allow the cell to make sure that various events have been checked off as completed before the next phase begins. Surveillance mechanisms that involve dozens of proteins, many just discovered, oversee these checkpoints. In the cell cycle, three checkpoints involve cyclin-dependent kinases; each has its own speci c cyclin that initiates either the G1, S, or mitosis phase. In addition, other checkpoints that don t involve cyclin-dependent kinases occur at other transition phases in the cell cycle. Cell cycle control is of particular interest because the cell cycle routinely halts if there is genetic damage, giving the cell a chance to repair the damage before committing to cell division. If the damage is too extreme, the cell can enter a programmed cell death sequence, discussed in chapter 16. If these mechanisms fail, cancer may result. The genetic control of the cell cycle is one of the most active areas of current research.
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Inactive complex P P
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Phosphates block active site
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Dephosphorylation P P
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Phosphorylation
G2 Cyclin B S Active complex M Active site G1 Breakdown of cyclin B Cdc2p
Inactive Cdc2p Buildup of cyclin B Figure 3.7 The proteins Cdc2p (CDK1) and cyclin B combine to form the maturation-promoting (or mitosis-promoting) factor. During mitosis, cyclin B is broken down. During G1 and S phases, cyclin B builds up and combines with Cdc2P, which is then phosphorylated at the active site to render it inactive. Dephosphorylation, a process that begins to take place only after DNA replication is nished, produces an active maturation-promoting factor.
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
3. Mitosis and Meiosis
The McGraw Hill Companies, 2001
Three
Mitosis and Meiosis
MITOSIS
Consider the engineering problem that mitosis must solve. Identical chromatids, called sister chromatids, the result of chromosomal replication, must separate so that each goes into a different daughter cell (see g. 3.3).These chromatids are the visible manifestation of the chromosomal replication that has taken place in the S phase of the cell cycle. The chromatids are initially held together; each will be called a chromosome when it separates and becomes independent. Each of the two daughter cells then ends up with a chromosome complement identical to that of the parent cell. Mitosis is nature s elegant process to achieve that end surely an engineering marvel. Mitosis is a continuous process. However, for descriptive purposes, we can break it into four stages: prophase, metaphase, anaphase, and telophase (Greek: pro-, before; meta-, mid; ana-, back; telo-, end). Replication (duplication) of the genetic material occurs during the S phase of the cell cycle (see g. 3.6). The timing of the four stages varies from species to species, from organ to organ within a species, and even from cell to cell within a given cell type.
Minus end
Tubulin
Plus end
Figure 3.8 Microtubules are hollow tubes made of and tubulin subunits that are constantly being added or removed.
Microtubules are formed from active centers called microtubule organizing centers. Centrioles, composed of two cylinders themselves composed of microtubules are microtubule organizing centers for cilia and agella. Under those circumstances, the centrioles are referred to as basal bodies. The centrioles were also originally believed to organize spindles. However, for most organisms, the microtubule organizing center is called the centrosome. In some organisms, such as fungi, a different cell organelle, the spindle pole body, serves this function. In most animals, the centrosome contains a centriole ( g. 3.9). However, the centriole is absent in most higher plants. Moreover, innovative ex-
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