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The Mitotic Spindle
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The process of mitosis involves an apparatus called the spindle. This structure is composed of microtubules, hollow cylinders made of protein subunits; each subunit is composed of one molecule of tubulin and one of tubulin; and each tubulin is the product of a different gene. (The spindle is named for the rounded rods, tapered at each end, once commonly used to hold yarn or thread.) Microtubules provide shape and structure to a eukaryotic cell as well as allow the cell to move its internal components and to move the cell itself with cilia and agella. Motion occurs as the microtubules slide past each other, a vesicle of some kind slides along the microtubules, and the microtubules shorten. Two proteins make up the microtubule motors that allow motion: kinesin and dynein. Scientists have studied microtubules through protein chemistry, through mutant organisms, and through innovative methods such as by coupling tubulin subunits with uorescing dyes to observe the microtubules in action. Microtubules are in a dynamic equilibrium, with subunits constantly being added or removed at both ends. On any microtubule, more activity occurs at one end than the other. The more active end of the tubule is called the plus end, the less active end the minus end ( g. 3.8). Both ends may be adding or removing subunits, or the plus end may be adding while the minus end is removing subunits. Generally, dynein causes movement toward the minus end, whereas kinesin causes movement toward the plus end of a microtubule, although exceptions exist.
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A centriole is composed of two barrels at right angles to each other. Each barrel is composed of nine tripartite units and a central cartwheel. Each of the three parts of a tripartite unit is a microtubule. Magni cation 111,800 . (Reproduced from The Journal of Cell Biology, 1968, Vol. 37, p. 381, by
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copyright permission of The Rockefeller University Press.)
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II. Mendelism and the Chromosomal Theory
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3. Mitosis and Meiosis
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Mitosis
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Centrosome Aster Interpolar microtubules Centriole + Nucleus Centrosome divides Nuclear membrane breaks down +
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Figure 3.10 Early in mitosis, the centrosome divides, and the separating halves move to opposite poles of the cell. This creates a
spindle in the middle of the cell after the nuclear membrane breaks down.
periments that removed the centrioles from cells that normally had them demonstrated that the centriole is not necessary for spindle formation. So, although we used to believe that the centriole formed the spindle in many organisms, we now know that the spindle is usually organized around the centrosome, which can function in this capacity without a centriole. The centriole, when present, replicates during the S and G2 phases. When mitosis begins, the centrosome divides and moves to opposite poles of the cell, around the nucleus ( g. 3.10). The centrosomes trail microtubules, forming the spindle, that at this point begin at each centrosome and overlap in the middle of the cell. These are called interpolar microtubules. Microtubules also spread out from the centrosome in the opposite direction from the spindle itself, forming an aster (see g. 3.10). The minus ends of microtubules emanate from the centrosome and the plus ends overlap in the middle of the cell. A third form of tubulin, tubulin, is needed to begin the formation of a microtubule.
chromosomes continue to shorten and thicken. The centromeres have already divided, and no new DNA synthesis is needed for the process to be completed. At this point, the sister chromatids are kept together by a complex, called cohesin, made up of at least four different protiens. Spindle bers are initially nucleated at the centrosome and grow outward into the cytoplasm ( g. 3.12). Some of these bers capture a kinetochore, the proteinaceous complex at the centromere of each sister chromatid; these bers are called kinetochore microtubules. At rst, one kinetochore or the other randomly attaches to a spindle ber. As the microtubules further move the chromosomes and as new microtubules attach and old microtubules break, each sister kinetochore eventually attaches to microtubules emanating from different poles. This ensures that sister chromatids move to opposite poles during anaphase. The number of microtubules that attach to each kinetochore differs in different species. It seems that 1 attaches to each kinetochore in yeast, 4 to 7 attach to each kinetochore in the cells of a rat fetus, and 70 to 150 attach in the plant Haemanthus (see g. 3.12).
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