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This stage of mitosis is characterized by the formation of the spindle and a shortening and thickening of the chromosomes so that individual chromosomes become visible. (We will discuss details of the molecular structure of the eukaryotic chromosome and the processes of coiling and shortening in chapter 15.) At this time also, the nuclear envelope (membrane) disintegrates and the nucleolus disappears ( g. 3.11). The nucleolus is a darkly stained body in the nucleus that is involved in ribosome construction and that forms around a nucleolar organizer locus on one of the chromosome pairs. The number of nucleoli varies in different species, but in the simplest case there are two nucleolar organizers per nucleus, one each on the two members of a homologous pair of chromosomes. Nucleoli re-form after mitosis. As prophase progresses, each chromosome is composed of two identical (sister) chromatids (see g. 3.3); the
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Interphase Early prophase Late prophase
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Nucleolus Nuclear membrane Figure 3.11 Nuclear events during interphase and prophase of
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mitosis. In this cell, 2n 4, consisting of one pair of long and one pair of short metacentric chromosomes. Maternal chromosomes are red; paternal chromosomes are blue. Note that each chromosome consists of two chromatids when the cell enters mitosis.
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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
moment, the cyclosome ubiquitinates the inhibitor, causing it to break down and freeing the separin to break down cohesin. This liberates the sister chromatids from each other (and is the instant when chromatids become chromosomes). The spindle then separates the sister chromatids in two stages, called anaphase A and anaphase B. In anaphase A, the chromosomes move toward the poles ( g. 3.14). During this process, the kinetochore itself acts as a microtubule motor, disassembling microtubules as it moves down them, pulling the chromosomes along ( g. 3.15). Thus, metacentric chromosomes appear V-shaped (as in g. 3.15), subtelocentrics appear J-shaped, and telocentrics appear rod-shaped. In anaphase B, the spindle itself elongates as overlapping interpolar microtubules slide apart. The general elongation of the spindle pulls the chromosomes apart.
Figure 3.12 Scanning electron micrograph of the centromeric
region of a metaphase chromosome from the plant Haemanthus katherinae. Spindle ber bundles on either side of the centromere extend in opposite directions. A ber not connected to the kinetochore is visible lying over the centromere. These bers are 60 to 70 nm in diameter. (Waheeb
K. Heneen, The centromeric region in the scanning electron microscope, Hereditas, 97 (1982): 311 14. Reproduced by permission.)
Telophase
At the end of anaphase ( g. 3.16), the separated sister chromatids (now full- edged chromosomes) have been pulled to opposite poles of the cell. The cell now reverses the steps of prophase to return to the interphase state ( g. 3.17).The chromosomes uncoil and begin to direct protein synthesis. A nuclear envelope re-forms around each set of chromosomes, nucleoli re-form, and cytokinesis takes place. The spindle breaks down into tubulin subunits; a residual of microtubules remains at
Metaphase
During metaphase, the chromosomes move to the equator of the cell. With the attachment of the spindle bers and the completion of the spindle itself, the chromosomes jockey into position in the equatorial plane of the spindle, called the metaphase plate. This happens as kinetochore microtubules exert opposing tension on the two sister kinetochores. Alignment of the chromosomes on this plate marks the end of metaphase ( g. 3.13).
Metaphase plate
Anaphase
During anaphase, the sister chromatids separate and move toward opposite poles on the spindle.The physical separation of the sister chromatids and their movement to opposite poles are two separate activities. Chromatid separation represents a checkpoint in the process of mitosis; a surveillance mechanism will not allow the process to continue until all chromosomes are lined up on the metaphase plate with their sister kinetochores held by microtubules from opposite poles. The surveillance mechanism somehow checks the physical tension the spindle bers exert on a pair of sister chromatids; an unpaired chromatid can delay or stop the process. Initially, an inhibitory protein called securin binds an enzyme called separin that can break down cohesin, the complex holding the chromatids together. At the correct
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