Mitosis 20369 225544768 2008-07-14T06:34:36Z زرشک 7201074 [[Image:300px-Anaphase-flourescent.jpg|thumb|right|[[Newt]] lung cell undergoing mitosis]] '''Mitosis''' is the process by which a [[eukaryotic]] cell separates the [[chromosome]]s in its [[cell nucleus]], into two identical sets in two daughter nuclei.<ref> Rubenstein, Irwin, and Susan M. Wick. "Cell." World Book Online Reference Center. 2008. 12 January 2008 <http://www.worldbookonline.com/wb/Article?id=ar102240> </ref> It is generally followed immediately by [[cytokinesis]], which divides the nuclei, [[cytoplasm]], [[organelle]]s and [[cell membrane]] into two daughter cells containing roughly equal shares of these cellular components. Mitosis and cytokinesis together define the '''mitotic (M) phase''' of the [[cell cycle]] - the [[cell division|division]] of the mother cell into two daughter cells, genetically identical to each other and to their parent cell. [[Image:Major events in mitosis.svg|right|thumb|350px|Mitosis divides the [[chromosome]]s in a [[cell nucleus]].]] Mitosis occurs exclusively in [[eukaryote|eukaryotic]] cells, but occurs in different ways in different species. For example, [[animal]]s undergo an "open" mitosis, where the [[nuclear envelope]] breaks down before the chromosomes separate, while [[fungi]] such as ''[[Aspergillus nidulans]]'' and ''[[Saccharomyces cerevisiae]]'' ([[yeast]]) undergo a "closed" mitosis, where chromosomes divide within an intact [[cell nucleus]].<ref>{{cite journal |author=De Souza CP, Osmani SA |title=Mitosis, not just open or closed |journal=Eukaryotic Cell |volume=6 |issue=9 |pages=1521–7 |year=2007 |pmid=17660363 |doi=10.1128/EC.00178-07}}</ref> [[Prokaryote|Prokaryotic]] cells, which lack a nucleus, divide by a process called [[binary fission]]. The process of mitosis is complex and highly regulated. The sequence of events is divided into phases, corresponding to the completion of one set of activities and the start of the next. These stages are prophase, prometaphase, metaphase, anaphase and telophase. During the process of mitosis the pairs of [[chromosomes]] condense and attach to fibers that pull the [[sister chromatids]] to opposite sides of the cell. The cell then divides in [[cytokinesis]], to produce two identical daughter cells.<ref>{{cite book | last = Maton | first = Anthea | authorlink = | coauthors = Hopkins, Jean Johnson, Susan LaHart, David, Quon Warner, David, Wright, Jill D | title = Cells: Building Blocks of Life | publisher = Prentice Hall | date = 1997 | location = New Jersey | pages = 70-74 | url = | doi = | id = | isbn = 0-13423476-6 }}</ref> Because cytokinesis usually occurs in conjunction with mitosis, "mitosis" is often used interchangeably with "mitotic phase". However, there are many cells where mitosis and cytokinesis occur separately, forming single cells with multiple nuclei. This occurs most notably among the [[fungus|fungi]] and [[slime mould]]s, but is found in various different groups. Even in animals, cytokinesis and mitosis may occur independently, for instance during certain stages of [[Drosophila melanogaster|fruit fly]] embryonic development.<ref name=Lilly>{{cite journal | author = Lilly M, Duronio R | title = New insights into cell cycle control from the Drosophila endocycle | journal = Oncogene | volume = 24 | issue = 17 | pages = 2765–75 | year = 2005 | pmid = 15838513 | doi = 10.1038/sj.onc.1208610 <!--Retrieved from CrossRef by DOI bot-->}}</ref> Errors in mitosis can either kill a cell through [[apoptosis]] or cause [[mutation]]s that may lead to [[cancer]]. ==Overview== The primary result of mitosis is the division of the parent cell's genome into two daughter cells. The genome is composed of a number of [[chromosome]]s, complexes of tightly-coiled [[DNA]] that contain [[DNA sequence|genetic information]] vital for proper cell function. Because each resultant daughter cell should be [[clone (genetics)|genetically identical]] to the parent cell, the parent cell must make a copy of each chromosome before mitosis. This occurs during S phase, in [[interphase]], the period that precedes the mitotic phase in the cell cycle where preparation for mitosis occurs.<ref name=Blow>{{cite journal | author = Blow J, Tanaka T | title = The chromosome cycle: coordinating replication and segregation. Second in the cycles review series | url=http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=16264427 | journal = EMBO Rep | volume = 6 | issue = 11 | pages = 1028–34 | year = 2005 | pmid = 16264427 | doi = 10.1038/sj.embor.7400557}}</ref> Each new chromosome now contains two identical copies of itself, called ''[[chromatid|sister chromatid]]s'', attached together in a specialized region of the chromosome known as the ''[[centromere]]''. Each sister chromatid is not considered a chromosome in itself, and a chromosome does not always contain two sister chromatids. In most [[eukaryotes]], the [[nuclear envelope]] that separates the DNA from the [[cytoplasm]] disassembles. The chromosomes align themselves in a line spanning the cell. [[Microtubule]]s, essentially miniature strings, splay out from opposite ends of the cell and shorten, pulling apart the sister chromatids of each chromosome.<ref>{{cite journal | author = Zhou J, Yao J, Joshi H | title = Attachment and tension in the spindle assembly checkpoint | journal = J Cell Sci | volume = 115 | issue = Pt 18 | pages = 3547–55 | year = 2002 | pmid = 12186941 | doi = 10.1242/jcs.00029 <!--Retrieved from CrossRef by DOI bot-->}}</ref> As a matter of convention, each sister chromatid is now considered a chromosome, so they are renamed to ''sister chromosomes''. As the cell elongates, corresponding sister chromosomes are pulled toward opposite ends. A new nuclear envelope forms around the separated sister chromosomes. As mitosis completes cytokinesis is well underway. In [[animal cell]]s, the cell pinches inward where the imaginary line used to be, (the pinching of the cell membrane to form the two daughter cells is called cleavage furrow) separating the two developing nuclei. In [[plant cell]]s, the daughter cells will construct a new dividing cell wall between each other. Eventually, the mother cell will be split in half, giving rise to two daughter cells, each with an equivalent and complete copy of the original genome. Prokaryotic cells undergo a process similar to mitosis called binary fission. However, prokaryotes cannot be properly said to undergo mitosis because they lack a nucleus and only have a single chromosome with no centromere.<ref>{{cite journal | author = Nanninga N | title = Cytokinesis in prokaryotes and eukaryotes: common principles and different solutions | url=http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=11381104 | journal = Microbiol Mol Biol Rev | volume = 65 | issue = 2 | pages = 319–33 | year = 2001 | pmid = 11381104 | doi = 10.1128/MMBR.65.2.319-333.2001 <!--Retrieved from CrossRef by DOI bot-->}}</ref> ==Mnemonic devices== The steps of mitosis can easily be remembered by a [[mnemonic device]] such as 'PMAT' or 'Please Make A Taco'. Each letter is the first letter of each step: 'P' for [[prophase]], 'M' for [[metaphase]], 'A' for [[anaphase]], and 'T' for [[telophase]]. Another mnemonic device for memorizing the steps of mitosis including [[interphase]] and [[cytokinesis]] is 'I Party More At The Club', or 'IPMATC'. Another way of remembering it is: 'In Paris Many Artists Teach': Interphase, Prophase, Metaphase, Anaphase, and Telophase. ==Phases of cell cycle and mitosis== ===Interphase=== {{main|Interphase}} [[Image:Cell cycle.png|thumb|right|The cell cycle]] <!-- that graph and the following paragraph needs to have full labels, not just single letters --> The mitotic phase is a relatively short period of the [[cell cycle]]. It alternates with the much longer ''[[interphase]]'', where the cell prepares itself for cell division. Interphase is divided into three phases, G<sub>1</sub> (first gap), S (synthesis), and G<sub>2</sub> (second gap). During all three phases, the cell grows by producing proteins and cytoplasmic organelles. However, chromosomes are replicated only during the [[S phase]]. Thus, a cell grows (G<sub>1</sub>), continues to grow as it duplicates its chromosomes (S), grows more and prepares for mitosis (G<sub>2</sub>), and divides (M).<ref name=Blow/> ===Preprophase=== {{main|Preprophase}} In plant cells only, prophase is preceded by a pre-prophase stage. In highly vacuolated plant cells, the nucleus has to migrate into the center of the cell before mitosis can begin. This is achieved through the formation of a [[phragmosome]], a transverse sheet of cytoplasm that bisects the cell along the future plane of cell division. In addition to phragmosome formation, preprophase is characterized by the formation of a ring of microtubules and actin filaments (called [[preprophase band]]) underneath the plasmamembrane around the equatorial plane of the future mitotic spindle and predicting the position of cell plate fusion during [[telophase]]. The cells of higher plants (such as the flowering plants) lack [[centrioles]]. Instead, spindle microtubules aggregate on the surface of the nuclear envelope during prophase. The preprophase band disappears during nuclear envelope disassembly and spindle formation in prometaphase.<ref>{{cite book | last = Raven | first = Peter H. | coauthors = Ray F. Evert, Susan E. Eichhorn | title = Biology of Plants, 7th Edition | publisher = W.H. Freeman and Company Publishers | date = 2005 | location = New York | pages = 58-67 | isbn = 0-7167-1007-2}}</ref> ===Prophase=== [[Image:Prophase.jpg|thumb|right|'''Prophase:''' The two round objects above the nucleus are the centrosomes. The chromatin has condensed.]] {{main|Prophase}} Normally, the genetic material in the nucleus is in a loosely bundled coil called [[chromatin]]. At the onset of prophase, chromatin condenses together into a highly ordered structure called a chromosome. Since the genetic material has already been duplicated earlier in S phase, the replicated chromosomes have two sister chromatids, bound together at the [[centromere]] by the cohesion complex. Chromosomes are visible at high magnification through a light microscope. Close to the nucleus are two [[centrosome]]s. Each centrosome, which was replicated earlier independent of mitosis, acts as a coordinating center for the cell's [[microtubule]]s. The two centrosomes nucleate microtubules (which may be thought of as cellular ropes or poles) by polymerizing soluble [[tubulin]] present in the cytoplasm. [[Molecular motor]] proteins create repulsive forces that will push the centrosomes to opposite side of the nucleus. The centrosomes are only present in animals. In plants the microtubules form independently. Some centrosomes contain a pair of [[centriole]]s that may help organize microtubule assembly, but they are not essential to formation of the mitotic spindle.<ref>{{cite journal | author = Lloyd C, Chan J. | title = Not so divided: the common basis of plant and animal cell division | journal = Nat Rev Mol Cell Biol. | volume = 7 | issue = 2 | pages = 147–52 | year = 2006 | pmid = 16493420 | doi = 10.1038/nrm1831 <!--Retrieved from CrossRef by DOI bot-->}}</ref> {{clear}} ===Prometaphase=== [[Image:Mitotic spindle color micrograph.gif|thumb|right|Micrograph showing condensed [[chromosome]]s in blue and the mitotic spindle in green during [[prometaphase]] of [[mitosis]] ]] [[Image:Prometaphase.jpg|thumb|right|'''Prometaphase:''' The nuclear membrane has degraded, and microtubules have invaded the nuclear space. These microtubules can attach to kinetochores or they can interact with opposing microtubules.]] {{main|Prometaphase}} The nuclear envelope disassembles and microtubules invade the nuclear space. This is called open mitosis, and it occurs in most multicellular organisms. Fungi and some [[protist]]s, such as [[algae]] or [[trichomonad]]s, undergo a variation called closed mitosis where the spindle forms inside the nucleus or its microtubules are able to penetrate an intact nuclear envelope.<ref>{{cite journal | author = Heywood P. | title = Ultrastructure of mitosis in the chloromonadophycean alga Vacuolaria virescens | journal = J Cell Sci. | volume = 31 | pages = 37–51 | year = 1978 | pmid = 670329}}</ref><ref>{{cite journal | author = Ribeiro K, Pereira-Neves A, Benchimol M | title = The mitotic spindle and associated membranes in the closed mitosis of trichomonads | journal = Biol Cell | volume = 94 | issue = 3 | pages = 157–72 | year = 2002 | pmid = 12206655 | doi = 10.1016/S0248-4900(02)01191-7 <!--Retrieved from CrossRef by DOI bot-->}}</ref> Each chromosome forms two [[kinetochore]]s at the centromere, one attached at each chromatid. A kinetochore is a complex protein structure that is analogous to a ring for the microtubule hook; it is the point where microtubules attach themselves to the chromosome.<ref>{{cite journal | author = Chan G, Liu S, Yen T | title = Kinetochore structure and function | journal = Trends Cell Biol | volume = 15 | issue = 11 | pages = 589–98 | year = 2005 | pmid = 16214339 | doi = 10.1016/j.tcb.2005.09.010 <!--Retrieved from CrossRef by DOI bot-->}}</ref> Although the kinetochore structure and function are not fully understood, it is known that it contains some form of [[List of gene families#Motor proteins|molecular motor]].<ref name=Maiato>{{cite journal | author = Maiato H, DeLuca J, Salmon E, Earnshaw W | title = The dynamic kinetochore-microtubule interface | doi= 10.1242/jcs.01536 | journal = J Cell Sci | volume = 117 | issue = Pt 23 | pages = 5461–77 | year = 2004 | pmid = 15509863}}</ref> When a microtubule connects with the kinetochore, the motor activates, using energy from [[Adenosine triphosphate|ATP]] to "crawl" up the tube toward the originating centrosome. This motor activity, coupled with polymerisation and depolymerisation of microtubules, provides the pulling force necessary to later separate the chromosome's two chromatids.<ref name=Maiato/> When the spindle grows to sufficient length, ''kinetochore microtubules'' begin searching for kinetochores to attach to. A number of ''nonkinetochore microtubules'' find and interact with corresponding nonkinetochore microtubules from the opposite centrosome to form the mitotic spindle.<ref name=Winey>{{cite journal | author = Winey M, Mamay C, O'Toole E, Mastronarde D, Giddings T, McDonald K, McIntosh J | title = Three-dimensional ultrastructural analysis of the Saccharomyces cerevisiae mitotic spindle | journal = J Cell Biol | volume = 129 | issue = 6 | pages = 1601–15 | year = 1995 | pmid = 7790357 | doi = 10.1083/jcb.129.6.1601 <!--Retrieved from CrossRef by DOI bot-->}}</ref> Prometaphase is sometimes considered part of prophase. {{clear}} ===Metaphase=== [[Image:Mitosis-flourescent.jpg|thumb|right|A cell in late [[metaphase]]. All chromosomes (blue) but one have arrived at the metaphase plate.]] [[Image:Metaphase.jpg|frame|right|'''Metaphase:''' The chromosomes have aligned at the metaphase plate.]] {{main|Metaphase}} As microtubules find and attach to kinetochores in prometaphase, the centromeres of the chromosomes convene along the ''metaphase plate'' or ''equatorial plane'', an imaginary line that is equidistant from the two centrosome poles.<ref name=Winey/> This even alignment is due to the counterbalance of the pulling powers generated by the opposing kinetochores, analogous to a tug-of-war between people of equal strength. In certain types of cells, chromosomes do not line up at the metaphase plate and instead move back and forth between the poles randomly, only roughly lining up along the midline. Metaphase comes from the [[Greek language|Greek]] ''μετα'' meaning "after." Because proper chromosome separation requires that every kinetochore be attached to a bundle of microtubules (spindle fibres) , it is thought that unattached kinetochores generate a signal to prevent premature progression to [[anaphase]][http://en.wikipedia.org/wiki/Mitosis#Anaphase] without all chromosomes being aligned. The signal creates the ''[[spindle checkpoint|mitotic spindle checkpoint]]''.<ref>{{cite journal | author = Chan G, Yen T | title = The mitotic checkpoint: a signaling pathway that allows a single unattached kinetochore to inhibit mitotic exit | journal = Prog Cell Cycle Res | volume = 5 | issue = | pages = 431–9 | year = | pmid = 14593737}}</ref> {{clear}} ===Anaphase=== [[Image:Anaphase.jpg|frame|right|'''Early anaphase:''' Kinetochore microtubules shorten]] {{main|Anaphase}} When every kinetochore is attached to a cluster of microtubules and the chromosomes have lined up along the metaphase plate, the cell proceeds to anaphase (from the [[Greek language|Greek]] ''ανα'' meaning “up,” “against,” “back,” or “re-”). Two events then occur; First, the proteins that bind sister chromatids together are cleaved, allowing them to separate. These sister chromatids turned sister chromosomes are pulled apart by shortening kinetochore microtubules and move toward the respective centrosomes to which they are attached. Next, the nonkinetochore microtubules elongate, pushing the centrosomes (and the set of chromosomes to which they are attached) apart to opposite ends of the cell. The force that causes the centrosomes to move towards the ends of the cell is still unknown, although there is a theory that suggests that the rapid assembly and breakdown of [[microtubules]] may cause this movement.<ref>{{cite journal | author = Kenneth R. Miller. | title = Anaphase | journal = Biology. | edition = 5 | pages = 169–170 | year = 2000}}</ref> These two stages are sometimes called early and late anaphase. Early anaphase is usually defined as the separation of the sister chromatids, while late anaphase is the elongation of the microtubules and the microtubules being pulled farther apart. At the end of anaphase, the cell has succeeded in separating identical copies of the genetic material into two distinct populations. {{clear}} ===Telophase=== [[Image:Telophase.jpg|frame|right|'''Telophase:''' The decondensing chromosomes are surrounded by nuclear membranes. Note cytokinesis has already begun, the pinching is known as the ''cleavage furrow''.]] {{main|Telophase}} Telophase (from the [[Greek language|Greek]] ''τελος'' meaning "end") is a reversal of prophase and prometaphase events. It "cleans up" the after effects of mitosis. At telophase, the nonkinetochore microtubules continue to lengthen, elongating the cell even more. Corresponding sister chromosomes attach at opposite ends of the cell. A new nuclear envelope, using fragments of the parent cell's nuclear membrane, forms around each set of separated sister chromosomes. Both sets of chromosomes, now surrounded by new nuclei, unfold back into chromatin. Mitosis is complete, but cell division is not yet complete. {{clear}} ===Cytokinesis=== {{main|Cytokinesis}} Cytokinesis is often mistakenly thought to be the final part of telophase, however cytokinesis is a separate process that begins at the same time as telophase. Cytokinesis is technically not even a phase of mitosis, but rather a separate process, necessary for completing cell division. In animal cells, a [[cleavage furrow]] (pinch) containing a contractile ring develops where the metaphase plate used to be, pinching off the separated nuclei.<ref>{{cite journal | author = Glotzer M | title = The molecular requirements for cytokinesis | journal = Science | volume = 307 | issue = 5716 | pages = 1735–9 | year = 2005 | pmid = 15774750 | doi = 10.1126/science.1096896 <!--Retrieved from CrossRef by DOI bot-->}}</ref> In both animal and plant cells, cell division is also driven by vesicles derived from the [[Golgi apparatus]], which move along microtubules to the middle of the cell.<ref>{{cite journal | author = Albertson R, Riggs B, Sullivan W | title = Membrane traffic: a driving force in cytokinesis | journal = Trends Cell Biol | volume = 15 | issue = 2 | pages = 92–101 | year = 2005 | pmid = 15695096 | doi = 10.1016/j.tcb.2004.12.008 <!--Retrieved from CrossRef by DOI bot-->}}</ref> In plants this structure coalesces into a cell plate at the center of the [[phragmoplast]] and develops into a cell wall, separating the two nuclei. The phragmoplast is a microtubule structure typical for higher plants, whereas some green algae use a [[phycoplast]] microtubule array during cytokinesis.<ref>{{cite book | last = Raven | first = Peter H. | coauthors = Ray F. Evert, Susan E. Eichhorn | title = Biology of Plants, 7th Edition | publisher = W.H. Freeman and Company Publishers | date = 2005 | location = New York | pages = 64-67, 328-329 | isbn = 0-7167-1007-2}}</ref> Each daughter cell has a complete copy of the genome of its parent cell. The end of cytokinesis marks the end of the M-phase. ==Significance== The importance of mitosis is the maintenance of the chromosomal set; each cell formed receives chromosomes that are alike in composition and equal in number to the chromosomes of the parent cell. Transcription is generally believed to cease during mitosis, but [[epigenetic]] mechanisms such as [[bookmarking]] function during this stage of the cell cycle to ensure that the "memory" of which genes were active prior to entry into mitosis are transmitted to the daughter cells.<ref>{{cite journal | author = Zhou G, Liu D, Liang C | title = Memory mechanisms of active transcription during cell division | journal = Bioessays | volume = 27 | issue = 12 | pages = 1239–45 | year = 2005 | pmid = 16299763 | doi = 10.1002/bies.20327 <!--Retrieved from CrossRef by DOI bot-->}}</ref> ==Consequences of errors== Although errors in mitosis are rare, the process may go wrong, especially during early cellular divisions in the [[zygote]]. Mitotic errors can be especially dangerous to the organism because future offspring from this parent cell will carry the same disorder. In ''non-disjunction'', a chromosome may fail to separate during anaphase. One daughter cell will receive both sister chromosomes and the other will receive none. This results in the former cell having three chromosomes coding for the same thing (two sisters and a homologue), a condition known as ''trisomy'', and the latter cell having only one chromosome (the homologous chromosome), a condition known as ''monosomy''. These cells are considered [[aneuploidy|aneuploidic]] cells and these abnormal cells can cause [[cancer]].<ref>{{cite journal | author = Draviam V, Xie S, Sorger P | title = Chromosome segregation and genomic stability | journal = Curr Opin Genet Dev | volume = 14 | issue = 2 | pages = 120–5 | year = 2004 | pmid = 15196457 | doi = 10.1016/j.gde.2004.02.007 <!--Retrieved from CrossRef by DOI bot-->}}</ref> Mitosis is a traumatic process. The cell goes through dramatic changes in ultrastructure, its organelles disintegrate and reform in a matter of hours, and chromosomes are jostled constantly by probing microtubules. Occasionally, chromosomes may become damaged. An arm of the chromosome may be broken and the fragment lost, causing [[genetic deletion|deletion]]. The fragment may incorrectly reattach to another, non-homologous chromosome, causing [[Chromosomal translocation|translocation]]. It may reattach to the original chromosome, but in reverse orientation, causing [[chromosomal inversion|inversion]]. Or, it may be treated erroneously as a separate chromosome, causing [[chromosomal duplication]]. The effect of these genetic abnormalities depend on the specific nature of the error. It may range from no noticeable effect, cancer induction, or organism death. ==Endomitosis== Endomitosis is a variant of mitosis without nuclear or cellular division, resulting in cells with many copies of the same chromosome occupying a single nucleus. This process may also be referred to as [[endoreduplication]] and the cells as [[Ploidy|endoploid]].<ref name=Lilly/> An example of a cell that goes through endomitosis is the [[megakaryocyte]].<ref name="pmid12871316">{{cite journal |author=Italiano JE, Shivdasani RA |title=Megakaryocytes and beyond: the birth of platelets |journal=J. Thromb. Haemost. |volume=1 |issue=6 |pages=1174–82 |year=2003 |pmid=12871316| doi = 10.1046/j.1538-7836.2003.00290.x <!--Retrieved from CrossRef by DOI bot-->}}</ref> ==Timeline in pictures== Real mitotic cells can be visualized through the microscope by [[staining (biology)|staining]] them with [[fluorescent]] antibodies and [[dyes]]. These light micrographs are included below. <gallery> <!-- Deleted image removed: Image:Prophase-flourescent.jpg|'''Early prophase:''' Nonkinetochore microtubules, shown as green strands, have established a matrix around the degrading nucleus, in blue. The green nodules are the centrosomes. --> Image:Prometaphase-flourescent.jpg|'''Early prometaphase:''' The nuclear membrane has just degraded, allowing the microtubules to quickly interact with the kinetochores on the chromosomes, which have just condensed. Image:Mitosis-flourescent.jpg|'''Late metaphase:''' The centrosomes have moved to the poles of the cell and have established the mitotic spindle. The chromosomes, in light blue, have all assembled at the metaphase plate, except for one. Image:Anaphase-flourescent.jpg|'''Anaphase:''' Lengthening nonkinetochore microtubules push the two sets of chromosomes further apart. </gallery> ==See also== *[[Meiosis]] *[[Cytoskeleton]] *[[Motor protein]] *[[Aneuploidy]] *[[Chromosomal disorder]] *[[Binary fission]] ==References== <div class="references-small" style="-moz-column-count:2; column-count:2;"> <references/> </div> ==Further reading== * Morgan DO (2007) "The Cell Cycle: Principles of Control" London: New Science Press. *{{cite web | author= Alberts B, Johnson A, Lewis J, Raff M, Roberts K, and Walter P | year= 2002 | url=http://www.ncbi.nlm.nih.gov/books/bv.fcgi?highlight=mitosis&rid=mboc4.section.3349 | title= Mitosis | format= | work= Molecular Biology of the Cell | publisher=Garland Science | accessdate = 2006-01-22}} * {{cite book | author = Campbell, N. and Reece, J. | date = December 2001 | chapter = The Cell Cycle | title = Biology | edition = 6th ed. | pages = pp. 217-224 | publisher = Benjamin Cummings/Addison-Wesley | location = San Francisco | id = ISBN 0-8053-6624-5 }} *{{cite web | author= Cooper, G. | year= 2000 | url=http://www.ncbi.nlm.nih.gov/books/bv.fcgi?highlight=M%20Phase,Events&rid=cooper.section.2470 | title= The Events of M Phase | format= | work= The Cell: A Molecular Approach | publisher=Sinaeur Associates, Inc | accessdate = 2006-01-22}} * {{cite book | last = Freeman | first = S | year = 2002 | chapter = Cell Division | title = Biological Science | pages = pp. 155-174 | publisher = Prentice Hall | location = Upper Saddle River, NJ | id = ISBN 0-13-081923-9 }} *{{cite web | author= Lodish H, Berk A, Zipursky L, Matsudaira P, Baltimore D, Darnell J | year= 2000 | url=http://www.ncbi.nlm.nih.gov/books/bv.fcgi?highlight=Overview,Control,Cell+Cycle&rid=mcb.section.3463 | title= Overview of the Cell Cycle and Its Control | format= | work= Molecular Cell Biology | publisher=W.H. Freeman | accessdate = 2006-01-22}} *{{cite book | last = Maton | first = Anthea | authorlink = | coauthors = Hopkins, Jean Johnson, Susan LaHart, David, Quon Warner, David, Wright, Jill D | title = Cells: Building Blocks of Life | publisher = Prentice Hall | date = 1997 | location = New Jersey | pages = 70-74 | url = | doi = | id = | isbn = 0-13423476-6 }} ==External links== * [http://www.scienceaid.co.uk/biology/genetics/mitosismeiosis.html Science aid: Mitosis and meiosis]: A simple account of the mitotic and meiotic processes. * [http://www.johnkyrk.com/mitosis.html Mitosis Animation]. * [http://iknow.net/player_window.html?movie=media/prophase_video.wmv&player=wm Video of a live amphibian lung cell undergoing mitosis]. * [http://www.pbs.org/wgbh/nova/miracle/divide.html# A Flash animation comparing Mitosis and Meiosis] *[http://www.cshprotocols.org/cgi/content/full/2007/3/pdb.prot4674 Studying Mitosis in Cultured Mammalian Cells] * [http://www.cellcycleontology.org CCO] The Cell-Cycle Ontology {{Cell cycle}} [[Category:Cell cycle|mitosis]] [[Category:Mitosis|*]] [[ar:انقسام فتيلي]] [[cs:Mitóza]] [[da:Mitose]] [[de:Mitose]] [[es:Mitosis]] [[eo:Mitozo]] [[fa:میتوز]] [[fr:Mitose]] [[gl:Mitose]] [[ko:유사 분열]] [[hr:Mitoza]] [[it:Mitosi]] [[he:מיטוזה]] [[lt:Mitozė]] [[hu:Mitózis]] [[mk:Митоза]] [[nl:Mitose]] [[ja:体細胞分裂]] [[no:Mitose]] [[pl:Mitoza]] [[pt:Mitose]] [[ru:Митоз]] [[simple:Mitosis]] [[sk:Mitóza]] [[sl:Mitoza]] [[sr:Митоза]] [[sv:Mitos]] [[th:ไมโทซิส]] [[uk:Мітоз]]