A Review of Mitosis: Eukaryotic Nuclear Division Cameron Incognito March 22, 2013 Introduction The ability of an organism to reproduce is one of the characteristics dividing living things from non-living things. Mitosis is a specific form of reproduction that occurs in eukaryotic, nongamete cells involving the division of nucleic genetic material into two identical daughter nuclei. Preceding the mitotic (M) phase is the interphase (consisting of the G1 phase, S phase, and the G2 phase) in which genetic material is duplicated and synthesized in preparation for the formation of daughter cells. Once interphase has been successfully completed, the cell progresses to the mitotic (M) phase, which is generally divided into five main stages: prophase, prometaphase, metaphase, anaphase, and telophase/cytokinesis. After mitosis and cytokinesis have been completed, a parent cell will have divided into two genetically identical daughter cells. The mitotic (M) phase is described in detail below in Figure 1: Figure 1. The five stages of the mitotic (M) phase Image from: < http://www.shmoop.com/cell-cycle/mitosis.html> Prophase Before the start of prophase, the nuclear envelope encases the nucleus, the chromosomes are loosely condensed, and two centrosomes have been developed in the G2 phase of interface. Prophase then begins starting with the chromatin fibers, a complex of DNA and proteins, becoming condensed into chromosomes. These chromosomes appear as two sister chromatids connected in the center at the centromere. The nucleoli begin to be degraded by cellular proteins causing the nucleoli to disappear. Lastly, the two centrosomes start to produce microtubules that 1|Page begin to extend, pushing the centrosomes towards opposite ends of the cell. The complex of centrosomes and microtubules is termed the mitotic spindle. Figure 2. A cell in prophase during the mitotic (M) phase of reproduction Image from: < http://www.biology.iupui.edu/biocourses/N100/2k4ch8mitosisnotes.html> Prometaphase During prometaphase, the nucleoli are completely degraded, releasing the chromosomes into the cytoplasm. The chromosomes continue to condense and form kinetochores at the centromere of chromatids. The microtubules expand through the cell entering into the nuclear space where specific microtubules, termed kinetochore microtubules, connect to the newly present kinetochores. The kinetochore microtubules are then moved around in a jerking motion attempting to move the chromatids towards the metaphase plate, a plane equidistant from each of the two centrosomes at the poles of the cell. Non-kinetochore microtubules begin to interact with complementary non-kinetochore microtubules being generated from the opposing centrosome. Figure 3. A cell in prometaphase during the mitotic (M) phase of reproduction Image from: < http://www.biology.iupui.edu/biocourses/N100/2k4ch8mitosisnotes.html> 2|Page Metaphase The centrosomes extend completely to opposite poles of the cell so the spindle is stretched as far as possible, providing ample space for the preparation of two cells. The chromosomes, connected to the spindle at the kinetochore, then reach equilibrium with their centromeres coming to rest at the metaphase plate. The kinetochores of sister chromatids are all fully attached to kinetochore microtubules branching from the opposite pole. Figure 4. A cell in metaphase during the mitotic (M) phase of reproduction Image from: < http://www.biology.iupui.edu/biocourses/N100/2k4ch8mitosisnotes.html> Anaphase At the beginning of anaphase, cohesion proteins are cleaved following a complex signaling cascade, allowing paired sister chromatids to split and form independent daughter chromosomes. The daughter chromosomes are pulled, centromere first, towards opposite poles of the cell due to the shortening of their kinetochore microtubules. The non-kinetochore microtubules continue to lengthen effectively elongating the cell. At the end of anaphase, each side of the cell contains equivalent and complete sets of chromosomes capable of forming a functioning cell. Figure 5. A cell in anaphase during the mitotic (M) phase of reproduction Image from: < http://www.biology.iupui.edu/biocourses/N100/2k4ch8mitosisnotes.html> 3|Page Telophase and Cytokinesis Telophase begins with the formation of two daughter nuclei in the cell. The nuclear envelope begins to form using the materials from the original parent nuclear envelope and the cell’s endomembrane system. The nucleoli begin to reappear inside of the envelope and the daughter chromosomes begin to de-condense inside of the nuclear space, transforming into accessible genetic information. The spindle microtubules, both kinetochore and non-kinetochore, depolymerize and are recycled in the cell. Mitosis, or the division of a single parent nucleus into two identical daughter nuclei, is complete after telophase. The cell, however, has not fully divided until cytokinesis, or the division of the cytoplasm, occurs. Cytokinesis begins with the development of a groove, termed the cleavage farrow, at the site of the metaphase plane. Actin microfilaments on one side of the groove interact with myosin proteins on the opposing side to cause a contraction until the cell is pinched in two. Two separate cells containing identical genetic information have now been produced. Figure 6. A cell after the completion of telophase and cytokinesis stages of reproduction Image from: < http://www.biology.iupui.edu/biocourses/N100/2k4ch8mitosisnotes.html> Conclusion Mitosis is the end stage of reproduction for all eukaryotic, non-gamete cells during which the cell’s genetic material is divided into two sets of identical daughter genetic information. With the successful completion of the interphase (G1 phase, S phase, and G2 phase), the cell proceeds to the mitotic (M) phase. Inside of the cell the nuclear envelope begins to degrade leaving the chromatin fibers exposed. The chromatin fibers tightly coil and condense producing chromosomes with a structure consisting of two identical sister chromatids joined at the centromere. The centrosomes start to produce the mitotic spindle and move away from each other towards opposite poles of the cell. Next, the nuclear envelope completely fragments allowing the extending kinetochore microtubules to attach at the chromatid centromeres. The 4|Page non-kinetochore microtubules continue to expand encompassing the entire cell until the centrosomes are at opposite poles of the cell. Once the chromosomes are aligned with the kinetochore on the metaphase plate, cohesion proteins are cleaved via a signaling cascade causing a transition to anaphase. During anaphase the two sister chromatids are pulled apart in opposite directions by kinetochore microtubules resulting in both sides of the cell containing equivalent and complete sets of genetic material. The non-kinetochore microtubules continue to expand elongating the cell. In telophase, two daughter nuclei are formed via the formation of new nuclear envelopes and the re-emergence of nucleoli. The microtubules are depolymerized, the chromosomes de-condense, and the cell proceeds to cytokinesis during which the cell divides its cytoplasm. The original parent cell has now been divided to create two identical daughter cells. References Reece, Jane B., Lisa A. Urry, Michael Lee Cain, Steven A. Wasserman, Peter V. Minorsky, and Robert Bradley Jackson. Campbell Biology. Boston [etc.: Benjamin Cummings, 2011. Print. 5|Page
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