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Introduction

Cell division is the process by which a parent cell divides into two or more daughter cells. Cell division usually occurs as part of a larger cell cycle. In eukaryotes, there are two distinct types of cell division; a vegetative division and the reproductive cell division. Both of these cell division cycles are used in the process of sexual reproduction at some point in their life cycle. Both are believed to be present in the last eukaryotic common ancestor. Prokaryotes (bacteria) undergo a vegetative cell division known as binary fission, where their genetic material is segregated equally into two daughter cells. Apart from binary fission, there are alternative manners of division such as budding. All cell divisions, regardless of organism, are preceded by a single round of DNA replication. For simple unicellular microorganisms such as the amoeba, one cell division is equivalent to reproduction – an entire new organism is created. On a larger scale, mitotic cell division can create progeny from multicellular organisms, such as plants that grow from cuttings. The primary concern of cell division is the maintenance of the original cell's genome. Before division can occur, the genomic information that is stored in chromosomes must be replicated, and the duplicated genome must be separated cleanly between cells.

Eukaryotes cell division

In eukaryotes there are two distinct types of cell division which are as followed:

1.     A vegetative division:

This is whereby each daughter cell is genetically identical to the parent cell. This process is called Mitosis. Mitotic cell division enables sexually reproducing organisms to develop from the one-celled zygote, which itself was produced by meiotic cell division from gametes. After growth, cell division by mitosis allows for continual construction and repair of the organism.

2.     A reproductive cell division:

This is where the number of chromosomes in the daughter cells is reduced by half to produce haploid gametes (meiosis). Meiosis results in four haploid daughter cells by undergoing one round of DNA replication followed by two divisions. Homologous chromosomes are separated in the first division, and sister chromatids are separated in the second division.

Phases of eukaryotic cell division

In eukaryotic cell division there are different stages in the cell cycle. These stages go from interphase, prophase, prometaphase, metaphase, anaphase, telophase and cytokinesis

Interphase

Interphase is the process a cell must go through before mitosis, meiosis, and cytokinesis. Interphase consists of three main phases: G1, S, and G2. G1 is a time of growth for the cell where specialized cellular functions occur in order to prepare the cell for DNA Replication. There are checkpoints during interphase that allow the cell to be either advance or halt further development. In S phase, the chromosomes are replicated in order for the genetic content to be maintained.During G2, the cell undergoes the final stages of growth before it enters the M phase, where spindles are synthesized. The M phase, can be either mitosis or meiosis depending on the type of cell. Germ cells, or gametes, undergo meiosis, while somatic cells will undergo mitosis. After the cell proceeds successfully through the M phase, it may then undergo cell division through cytokinesis. The control of each checkpoint is controlled by cyclin and cyclin-dependent kinases. The progression of interphase is the result of the increased amount of cyclin. As the amount of cyclin increases, more and more cyclin dependent kinases attach to cyclin signaling the cell further into interphase. At the peak of the cyclin, attached to the cyclin dependent kinases this system pushes the cell out of interphase and into the M phase, where mitosis, meiosis, and cytokinesis occur. There are three transition checkpoints the cell has to go through before entering the M phase. The most important being the G1-S transition checkpoint. If the cell does not pass this checkpoint, it results in the cell exiting the cell cycle.

Prophase

Prophase is the first stage of division. The nuclear envelope is broken down, long strands of chromatin condense to form shorter more visible strands called chromosomes, the nucleolus disappears, and microtubules attach to the chromosomes at the kinetochores present in the centromere. Microtubules associated with the alignment and separation of chromosomes are referred to as the spindle and spindle fibers. Chromosomes will also be visible under a microscope and will be connected at the centromere. During this condensation and alignment period in meiosis, the homologous chromosomes undergo a break in their double-stranded DNA at the same locations, followed by a recombination of the now fragmented parental DNA strands into non-parental combinations, known as crossing over. This process is evidenced to be caused in a large part by the highly conserved Spo11 protein through a mechanism similar to that seen with toposomerase in DNA replication and transcription.


Prometaphase

Prometaphase is the second stage or phase of mitosis. In this stage the duplicated material in the nucleus is separated into two identical daughter cells. One important event that takes place is the nuclear envelop breaks down resulting into the freedom of the sister chromatids. This happens to separate the two daughter cells. In addition, during this phase, a protein named Kinetochore is formed around the centromere. Protein filaments named kinetochore microtubules extend from poles on both sides of the cell and attach themselves to the kinetochores. Once this phase has been successfully completed, the cell moves onto the third phase called metaphase

Metaphase

In metaphase, the centromeres of the chromosomes convene themselves on the metaphase plate (or equatorial plate), an imaginary line that is equidistant from the two centrosome poles and held together by complex complexes known as cohesins. Chromosomes line up in the middle of the cell by microtubule organizing centers (MTOCs) pushing and pulling on centromeres of both chromatids thereby causing the chromosome to move to the center. At this point the chromosomes are still condensing and are currently one step away from being the most coiled and condensed they will be, and the spindle fibers have already connected to the kinetochores. During this phase all the microtubules, with the exception of the kinetochores, are in a state of instability promoting their progression towards anaphase. At this point, the chromosomes are ready to split into opposite poles of the cell towards the spindle to which they are connected.

Anaphase

Anaphase is a very short stage of the cell cycle and occurs after the chromosomes align at the mitotic plate. Kinetochores emit anaphase-inhibition signals until their attachment to the mitotic spindle. Once the final chromosome is properly aligned and attached the final signal dissipates and triggers the abrupt shift to anaphase. This abrupt shift is caused by the activation of the anaphase-promoting complex and its function of tagging degradation of proteins important towards the metaphase-anaphase transition. One of these proteins that is broken down is securin which through its breakdown releases the enzyme separase that cleaves the cohesin rings holding together the sister chromatids thereby leading to the chromosomes separating. After the chromosomes line up in the middle of the cell, the spindle fibers will pull them apart. The chromosomes are split apart while the sister chromatids move to opposite sides of the cell. As the sister chromatids are being pulled apart, the cell and plasma are elongated by non-kinetochore microtubules.

Telophase

Telophase is one of the last stages of the cell cycle in which a cleavage furrow splits the cells cytoplasm (cytokinesis) and chromatin. This occurs through the synthesis of a new nuclear envelopes that forms around the chromatin gathered at each pole and the reformation of the nucleolus as the chromosomes decondense their chromatin back to the loose state it possessed during interphase. The division of the cellular contents is not always equal and can vary by cell type as seen with oocyte formation where one of the four daughter cells possess the majority of the cytoplasm.


Variants

Cells are broadly classified into two main categories:

1.     Prokaryotic cells: These are simple, single-celled, non-nucleated cells that lack membrane-bound organelles.

2.      Eukaryotic cells: These are complex, nucleated cells that contain organelles. They are also enclosed by a plasma membrane.

These differences result in the divisions of eukaryotic and prokaryotic cells being different. Due to their structural differences, eukaryotic and prokaryotic cells do not divide in the same way. Also, the pattern of cell division that transforms eukaryotic stem cells into gametes (sperm cells in males or egg cells in females), termed meiosis, is different from that of the division of somatic cells in the body.


Degradation

Multicellular organisms replace worn-out cells through cell division. In some animals, however, cell division eventually halts. In humans this occurs, on average, after 52 divisions, known as the Hayflick limit. The cell is then referred to as senescent. With each division the cells telomeres, protective sequences of DNA on the end of a chromosome that prevent degradation of the chromosomal DNA, shorten. This shortening has been correlated to negative effects such as age related diseases and shortened lifespans in humans. Cancer cells, on the other hand, are not thought to degrade in this way, if at all. An enzyme complex called telomerase, present in large quantities in cancerous cells, rebuilds the telomeres through synthesis of telomeric DNA repeats, allowing division to continue indefinitely.

History

A cell division under microscope was first discovered by German botanist Hugo von Mohl in 1835 as he worked over the green alga Cladophoraglomerata.

In 1943, cell division was filmed for the first time by Kurt Michel using a phase-contrast microscope.