Mitosis

In cell biology, mitosis (/maɪˈtoʊsɪs/) is a part of the cell cycle in which replicated chromosomes
are separated into two new nuclei. Cell division gives rise to genetically identical cells in which
the total number of chromosomes is maintained.[1] Therefore, mitosis is also known as
equational division.[2][3] In general, mitosis is preceded by S phase of interphase (during which
DNA replication occurs) and is often followed by telophase and cytokinesis; which divides the
cytoplasm, organelles and cell membrane of one cell into two new cells containing roughly equal
shares of these cellular components.[4] The different stages of mitosis altogether define the
mitotic (M) phase of an animal cell cycle—the division of the mother cell into two daughter cells
genetically identical to each other.[5]
Mitosis in an animal cell (phases ordered counter-clockwise).

Mitosis divides the chromosomes in a cell nucleus.

Label-free live cell imaging of Mesenchymal Stem Cells undergoing mitosis

Onion (Allium) cells in different phases of the cell cycle enlarged 800 diameters.

a. non-dividing cells

b. nuclei preparing for division (spireme-stage)

c. dividing cells showing mitotic figures

e. pair of daughter-cells shortly after division

The process of mitosis is divided into stages corresponding to the completion of one set of
activities and the start of the next. These stages are preprophase (specific to plant cells),
prophase, prometaphase, metaphase, anaphase, and telophase. During mitosis, the
chromosomes, which have already duplicated, condense and attach to spindle fibers that pull
one copy of each chromosome to opposite sides of the cell.[6] The result is two genetically
identical daughter nuclei. The rest of the cell may then continue to divide by cytokinesis to
produce two daughter cells.[7] The different phases of mitosis can be visualized in real time,
using live cell imaging.[8] Producing three or more daughter cells instead of the normal two is a
mitotic error called tripolar mitosis or multipolar mitosis (direct cell triplication /
multiplication).[9] Other errors during mitosis can induce apoptosis (programmed cell death) or
cause mutations. Certain types of cancer can arise from such mutations.[10]

Mitosis occurs only in eukaryotic cells. Prokaryotic cells, which lack a nucleus, divide by a
different process called binary fission. Mitosis varies between organisms.[11] For example,
animal cells undergo an "open" mitosis, where the nuclear envelope breaks down before the
chromosomes separate, whereas fungi undergo a "closed" mitosis, where chromosomes divide
within an intact cell nucleus.[12] Most animal cells undergo a shape change, known as mitotic
cell rounding, to adopt a near spherical morphology at the start of mitosis. Most human cells are
produced by mitotic cell division. Important exceptions include the gametes – sperm and egg
cells – which are produced by meiosis.

Discovery

Numerous descriptions of cell division were made during 18th and 19th centuries, with various
degrees of accuracy.[13] In 1835, the German botanist Hugo von Mohl, described cell division in
the green algae Cladophora glomerata, stating that multiplication of cells occurs through cell
division.[14][15][16] In 1838, Matthias Jakob Schleiden affirmed that "formation of new cells in their
interior was a general rule for cell multiplication in plants", a view later rejected in favour of
Mohl's model, due to contributions of Robert Remak and others.[17]

In animal cells, cell division with mitosis was discovered in frog, rabbit, and cat cornea cells in
1873 and described for the first time by the Polish histologist Wacław Mayzel in 1875.[18][19]

Bütschli, Schneider and Fol might have also claimed the discovery of the process presently
known as "mitosis".[13] In 1873, the German zoologist Otto Bütschli published data from
observations on nematodes. A few years later, he discovered and described mitosis based on
those observations.[20][21][22]

The term "mitosis", coined by Walther Flemming in 1882,[23] is derived from the Greek word
μίτος (mitos, "warp thread").[24][25] There are some alternative names for the process,[26] e.g.,
"karyokinesis" (nuclear division), a term introduced by Schleicher in 1878,[27][28] or "equational
division", proposed by August Weismann in 1887.[29] However, the term "mitosis" is also used in
a broad sense by some authors to refer to karyokinesis and cytokinesis together.[30] Presently,
"equational division" is more commonly used to refer to meiosis II, the part of meiosis most like
mitosis.[31]

Phases

Overview

Play media

Time-lapse video of mitosis in a Drosophila melanogaster embryo

The primary result of mitosis and cytokinesis is the transfer of a parent cell's genome into two
daughter cells. The genome is composed of a number of chromosomes—complexes of tightly
coiled DNA that contain genetic information vital for proper cell function.[32] Because each
resultant daughter cell should be genetically identical to the parent cell, the parent cell must
make a copy of each chromosome before mitosis. This occurs during the S phase of
interphase.[33] Chromosome duplication results in two identical sister chromatids bound together
by cohesin proteins at the centromere.

When mitosis begins, the chromosomes condense and become visible. In some eukaryotes, for
example animals, the nuclear envelope, which segregates the DNA from the cytoplasm,
disintegrates into small vesicles. The nucleolus, which makes ribosomes in the cell, also
disappears. Microtubules project from opposite ends of the cell, attach to the centromeres, and
align the chromosomes centrally within the cell. The microtubules then contract to pull the sister
chromatids of each chromosome apart.[34] Sister chromatids at this point are called daughter
chromosomes. As the cell elongates, corresponding daughter chromosomes are pulled toward
opposite ends of the cell and condense maximally in late anaphase. A new nuclear envelope
forms around the separated daughter chromosomes, which decondense to form interphase
nuclei.

During mitotic progression, typically after the anaphase onset, the cell may undergo cytokinesis.
In animal cells, a cell membrane pinches inward between the two developing nuclei to produce
two new cells. In plant cells, a cell plate forms between the two nuclei. Cytokinesis does not
always occur; coenocytic (a type of multinucleate condition) cells undergo mitosis without
cytokinesis.

Diagram of the mitotic phases

Interphase

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 the process of cell division. Interphase is divided
into three phases: G1 (first gap), S (synthesis), and G2 (second gap). During all three parts of
interphase, the cell grows by producing proteins and cytoplasmic organelles. However,
chromosomes are replicated only during the S phase. Thus, a cell grows (G1), continues to grow
as it duplicates its chromosomes (S), grows more and prepares for mitosis (G2), and finally
divides (M) before restarting the cycle.[33] All these phases in the cell cycle are highly regulated
by cyclins, cyclin-dependent kinases, and other cell cycle proteins. The phases follow one
another in strict order and there are "checkpoints" that give the cell cues to proceed from one
phase to another.[35] Cells may also temporarily or permanently leave the cell cycle and enter G0
phase to stop dividing. This can occur when cells become overcrowded (density-dependent
inhibition) or when they differentiate to carry out specific functions for the organism, as is the
case for human heart muscle cells and neurons. Some G0 cells have the ability to re-enter the
cell cycle.

DNA double-strand breaks can be repaired during interphase by two principal processes.[36] The
first process, non-homologous end joining (NHEJ), can join the two broken ends of DNA in the
G1, S and G2 phases of interphase. The second process, homologous recombinational repair
(HRR), is more accurate than NHEJ in repairing double-strand breaks. HRR is active during the S
and G2 phases of interphase when DNA replication is either partially accomplished or after it is
completed, since HRR requires two adjacent homologs.

Interphase helps prepare the cell for mitotic division. It dictates whether the mitotic cell division
will occur. It carefully stops the cell from proceeding whenever the cell's DNA is damaged or has
not completed an important phase. The interphase is very important as it will determine if
mitosis completes successfully. It will reduce the amount of damaged cells produced and the
production of cancerous cells. A miscalculation by the key Interphase proteins could be crucial
as the latter could potentially create cancerous cells.[37] Today, more research is being done to
understand specifically how the phases stated above occur.

Mitosis
Stages of early mitosis in a vertebrate cell with micrographs of chromatids

Preprophase (plant cells)

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 plasma membrane around the equatorial plane of the
future mitotic spindle. This band marks the position where the cell will eventually divide. The
cells of higher plants (such as the flowering plants) lack centrioles; instead, microtubules form a
spindle on the surface of the nucleus and are then organized into a spindle by the chromosomes
themselves, after the nuclear envelope breaks down.[38] The preprophase band disappears
during nuclear envelope breakdown and spindle formation in prometaphase.[39]: 58–67 
Prophase

Condensing chromosomes. Interphase nucleus (left), condensing chromosomes (middle) and condensed chromosomes
(right).

Prophase during mitosis

During prophase, which occurs after G2 interphase, the cell prepares to divide by tightly
condensing its chromosomes and initiating mitotic spindle formation. During interphase, the
genetic material in the nucleus consists of loosely packed chromatin. At the onset of prophase,
chromatin fibers condense into discrete chromosomes that are typically visible at high
magnification through a light microscope. In this stage, chromosomes are long, thin, and thread-
like. Each chromosome has two chromatids. The two chromatids are joined at the centromere.

Gene transcription ceases during prophase and does not resume until late anaphase to early G1
phase.[40][41][42] The nucleolus also disappears during early prophase.[43]

Close to the nucleus of animal cells are structures called centrosomes, consisting of a pair of
centrioles surrounded by a loose collection of proteins. The centrosome is the coordinating
center for the cell's microtubules. A cell inherits a single centrosome at cell division, which is
duplicated by the cell before a new round of mitosis begins, giving a pair of centrosomes. The
two centrosomes polymerize tubulin to help form a microtubule spindle apparatus. Motor
proteins then push the centrosomes along these microtubules to opposite sides of the cell.
Although centrosomes help organize microtubule assembly, they are not essential for the
formation of the spindle apparatus, since they are absent from plants,[38] and are not absolutely
required for animal cell mitosis.[44]

Prometaphase

At the beginning of prometaphase in animal cells, phosphorylation of nuclear lamins causes the
nuclear envelope to disintegrate into small membrane vesicles. As this happens, microtubules
invade the nuclear space. This is called open mitosis, and it occurs in some multicellular
organisms. Fungi and some protists, such as algae or trichomonads, undergo a variation called
closed mitosis where the spindle forms inside the nucleus, or the microtubules penetrate the
intact nuclear envelope.[45][46]

In late prometaphase, kinetochore microtubules begin to search for and attach to chromosomal
kinetochores.[47] A kinetochore is a proteinaceous microtubule-binding structure that forms on
the chromosomal centromere during late prophase.[47][48] A number of polar microtubules find
and interact with corresponding polar microtubules from the opposite centrosome to form the
mitotic spindle.[49] Although the kinetochore structure and function are not fully understood, it is
known that it contains some form of molecular motor.[50] When a microtubule connects with the
kinetochore, the motor activates, using energy from 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.[50]

Metaphase
A cell in late metaphase. All chromosomes (blue) but one have arrived at the metaphase plate.

Metaphase during Mitosis

After the microtubules have located and attached to the kinetochores in prometaphase, the two
centrosomes begin pulling the chromosomes towards opposite ends of the cell. The resulting
tension causes the chromosomes to align along the metaphase plate or equatorial plane, an
imaginary line that is centrally located between the two centrosomes (at approximately the
midline of the cell).[49] To ensure equitable distribution of chromosomes at the end of mitosis,
the metaphase checkpoint guarantees that kinetochores are properly attached to the mitotic
spindle and that the chromosomes are aligned along the metaphase plate.[51] If the cell
successfully passes through the metaphase checkpoint, it proceeds to anaphase.

Anaphase
Anaphase during Mitosis

During anaphase A, the cohesins that bind sister chromatids together are cleaved, forming two
identical daughter chromosomes.[52] Shortening of the kinetochore microtubules pulls the newly
formed daughter chromosomes to opposite ends of the cell. During anaphase B, polar
microtubules push against each other, causing the cell to elongate.[53] In late anaphase,
chromosomes also reach their overall maximal condensation level, to help chromosome
segregation and the re-formation of the nucleus.[54] In most animal cells, anaphase A precedes
anaphase B, but some vertebrate egg cells demonstrate the opposite order of events.[52]

Telophase

Telophase during mitosis

Telophase (from the Greek word τελος meaning "end") is a reversal of prophase and
prometaphase events. At telophase, the polar microtubules continue to lengthen, elongating the
cell even more. If the nuclear envelope has broken down, a new nuclear envelope forms using
the membrane vesicles of the parent cell's old nuclear envelope. The new envelope forms
around each set of separated daughter chromosomes (though the membrane does not enclose
the centrosomes) and the nucleolus reappears. Both sets of chromosomes, now surrounded by
new nuclear membrane, begin to "relax" or decondense. Mitosis is complete. Each daughter
nucleus has an identical set of chromosomes. Cell division may or may not occur at this time
depending on the organism.

Cytokinesis

Cytokinesis illustration

Cilliate undergoing cytokinesis, with the cleavage furrow being clearly visible
Cytokinesis is not 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.[55] 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.[56] 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.[39]: 64–7, 328–9  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.

There are many cells where mitosis and cytokinesis occur separately, forming single cells with
multiple nuclei. The most notable occurrence of this is among the fungi, slime molds, and
coenocytic algae, but the phenomenon is found in various other organisms. Even in animals,
cytokinesis and mitosis may occur independently, for instance during certain stages of fruit fly
embryonic development.[57]

Function

Mitosis's "function" or significance relies on the maintenance of the chromosomal set; each
formed cell receives chromosomes that are alike in composition and equal in number to the
chromosomes of the parent cell.

Mitosis occurs in the following circumstances:

Development and growth: The number of cells within an organism increases by mitosis. This
is the basis of the development of a multicellular body from a single cell, i.e., zygote and also
the basis of the growth of a multicellular body.

Cell replacement: In some parts of the body, e.g. skin and digestive tract, cells are constantly
sloughed off and replaced by new ones. New cells are formed by mitosis and so are exact
copies of the cells being replaced. In like manner, red blood cells have a short lifespan (only
about 4 months) and new RBCs are formed by mitosis.

Regeneration: Some organisms can regenerate body parts. The production of new cells in
such instances is achieved by mitosis. For example, starfish regenerate lost arms through
mitosis.
 Asexual reproduction: Some organisms produce genetically similar offspring through asexual
reproduction. For example, the hydra reproduces asexually by budding. The cells at the
surface of hydra undergo mitosis and form a mass called a bud. Mitosis continues in the cells
of the bud and this grows into a new individual. The same division happens during asexual
reproduction or vegetative propagation in plants.

Variations

Forms of mitosis

The mitosis process in the cells of eukaryotic organisms follows a similar pattern, but with
variations in three main details. "Closed" and "open" mitosis can be distinguished on the basis of
nuclear envelope remaining intact or breaking down. An intermediate form with partial
degradation of the nuclear envelope is called "semiopen" mitosis. With respect to the symmetry
of the spindle apparatus during metaphase, an approximately axially symmetric (centered)
shape is called "orthomitosis", distinguished from the eccentric spindles of "pleuromitosis", in
which mitotic apparatus has bilateral symmetry. Finally, a third criterion is the location of the
central spindle in case of closed pleuromitosis: "extranuclear" (spindle located in the cytoplasm)
or "intranuclear" (in the nucleus).[11]
 closed
intranuclear

pleuromitosis

closed
extranuclear

pleuromitosis

closed
orthomitosis
 semiopen

pleuromitosis

semiopen

orthomitosis

open

orthomitosis

Nuclear division takes place only in cells of organisms of the eukaryotic domain, as bacteria and
archaea have no nucleus. Bacteria and archaea undergo a different type of division.Within each
of the eukaryotic supergroups, mitosis of the open form can be found, as well as closed mitosis,
except for Excavata, which show exclusively closed mitosis.[58] Following, the occurrence of the
forms of mitosis in eukaryotes:[11][59]
 Closed intranuclear pleuromitosis is typical of Foraminifera, some Prasinomonadida, some
Kinetoplastida, the Oxymonadida, the Haplosporidia, many fungi (chytrids, oomycetes,
zygomycetes, ascomycetes), and some Radiolaria (Spumellaria and Acantharia); it seems to
be the most primitive type.

Closed extranuclear pleuromitosis occurs in Trichomonadida and Dinoflagellata.

Closed orthomitosis is found among diatoms, ciliates, some Microsporidia, unicellular yeasts
and some multicellular fungi.

Semiopen pleuromitosis is typical of most Apicomplexa.

Semiopen orthomitosis occurs with different variants in some amoebae (Lobosa) and some
green flagellates (e.g., Raphidophyta or Volvox).

Open orthomitosis is typical in mammals and other Metazoa, and in land plants; but it also
occurs in some protists.

Errors and other variations

An abnormal (tripolar) mitosis (12 o'clock position) in a precancerous lesion of the stomach (H&E stain)

Errors can occur during mitosis, especially during early embryonic development in humans.[60]
During each step of mitosis, there are normally checkpoints as well that control the normal
outcome of mitosis.[61] But, occasionally to almost rarely, mistakes will happen. Mitotic errors
can create aneuploid cells that have too few or too many of one or more chromosomes, a
condition associated with cancer.[62][63] Early human embryos, cancer cells, infected or
intoxicated cells can also suffer from pathological division into three or more daughter cells
(tripolar or multipolar mitosis), resulting in severe errors in their chromosomal complements.[9]
In nondisjunction, sister chromatids fail to separate during anaphase.[64] One daughter cell
receives both sister chromatids from the nondisjoining chromosome and the other cell receives
none. As a result, the former cell gets three copies of the chromosome, a condition known as
trisomy, and the latter will have only one copy, a condition known as monosomy. On occasion,
when cells experience nondisjunction, they fail to complete cytokinesis and retain both nuclei in
one cell, resulting in binucleated cells.[65]

Anaphase lag occurs when the movement of one chromatid is impeded during anaphase.[64] This
may be caused by a failure of the mitotic spindle to properly attach to the chromosome. The
lagging chromatid is excluded from both nuclei and is lost. Therefore, one of the daughter cells
will be monosomic for that chromosome.

Endoreduplication (or endoreplication) occurs when chromosomes duplicate but the cell does
not subsequently divide. This results in polyploid cells or, if the chromosomes duplicates
repeatedly, polytene chromosomes.[64][66] Endoreduplication is found in many species and
appears to be a normal part of development.[66] Endomitosis is a variant of endoreduplication in
which cells replicate their chromosomes during S phase and enter, but prematurely terminate,
mitosis. Instead of being divided into two new daughter nuclei, the replicated chromosomes are
retained within the original nucleus.[57][67] The cells then re-enter G1 and S phase and replicate
their chromosomes again.[67] This may occur multiple times, increasing the chromosome
number with each round of replication and endomitosis. Platelet-producing megakaryocytes go
through endomitosis during cell differentiation.[68][69]

Amitosis in ciliates and in animal placental tissues results in a random distribution of parental
alleles.

Karyokinesis without cytokinesis originates multinucleated cells called coenocytes.

Diagnostic marker
Mitosis appearances in breast cancer

In histopathology, the mitosis rate (mitotic count or mitotic index) is an important parameter in
various types of tissue samples, for diagnosis as well as to further specify the aggressiveness
of tumors. For example, there is routinely a quantification of mitotic count in breast cancer
classification.[70] The mitoses must be counted in an area of the highest mitotic activity. Visually
identifying these areas, is difficult in tumors with very high mitotic activity.[71] Also, the detection
of atypical forms of mitosis can be used both as a diagnostic and prognostic marker. For
example, lag-type mitosis (non-attached condensed chromatin in the area of the mitotic figure)
indicates high risk human papillomavirus infection-related Cervical cancer. In order to improve
the reproducibilty and accuracy of the mitotic count, automated image analysis using deep
learning-based algorithms have been proposed.[72] However, further research is needed before
those algorithms can be used to routine diagnostics.
 Normal and atypical forms of mitosis in cancer cells. A, normal mitosis; B, chromatin bridge; C,
multipolar mitosis; D, ring mitosis; E, dispersed mitosis; F, asymmetrical mitosis; G, lag-type mitosis;
and H, micronuclei. H&E stain.

Related cell processes

Cell rounding
Cell shape changes through mitosis for a typical animal cell cultured on a flat surface. The cell undergoes mitotic cell
rounding during spindle assembly and then divides via cytokinesis. The actomyosin cortex is depicted in red,
DNA/chromosomes purple, microtubules green, and membrane and retraction fibers in black. Rounding also occurs in live
tissue, as described in the text.

In animal tissue, most cells round up to a near-spherical shape during mitosis.[73][74][75] In

epithelia and epidermis, an efficient rounding process is correlated with proper mitotic spindle
alignment and subsequent correct positioning of daughter cells.[74][75][76][77] Moreover,
researchers have found that if rounding is heavily suppressed it may result in spindle defects,
primarily pole splitting and failure to efficiently capture chromosomes.[78] Therefore, mitotic cell
rounding is thought to play a protective role in ensuring accurate mitosis.[77][79]

Rounding forces are driven by reorganization of F-actin and myosin (actomyosin) into a
contractile homogeneous cell cortex that 1) rigidifies the cell periphery[79][80][81] and 2) facilitates
generation of intracellular hydrostatic pressure (up to 10 fold higher than interphase).[82][83][84]
The generation of intracellular pressure is particularly critical under confinement, such as would
be important in a tissue scenario, where outward forces must be produced to round up against
surrounding cells and/or the extracellular matrix. Generation of pressure is dependent on formin-
mediated F-actin nucleation[84] and Rho kinase (ROCK)-mediated myosin II contraction,[80][82][84]
both of which are governed upstream by signaling pathways RhoA and ECT2[80][81] through the
activity of Cdk1.[84] Due to its importance in mitosis, the molecular components and dynamics of
the mitotic actomyosin cortex is an area of active research.

Mitotic recombination

Mitotic cells irradiated with X-rays in the G1 phase of the cell cycle repair recombinogenic DNA
damages primarily by recombination between homologous chromosomes.[85] Mitotic cells
irradiated in the G2 phase repair such damages preferentially by sister-chromatid
recombination.[85] Mutations in genes encoding enzymes employed in recombination cause cells
to have increased sensitivity to being killed by a variety of DNA damaging agents.[86][87][88] These
findings suggest that mitotic recombination is an adaptation for repairing DNA damages
including those that are potentially lethal.

Evolution
Some types of cell division in prokaryotes and eukaryotes

There are prokaryotic homologs of all the key molecules of eukaryotic mitosis (e.g., actins,
tubulins). Being a universal eukaryotic property, mitosis probably arose at the base of the
eukaryotic tree. As mitosis is less complex than meiosis, meiosis may have arisen after
mitosis.[89] However, sexual reproduction involving meiosis is also a primitive characteristic of
eukaryotes.[90] Thus meiosis and mitosis may both have evolved, in parallel, from ancestral
prokaryotic processes.

While in bacterial cell division, after duplication of DNA, two circular chromosomes are attached
to a special region of the cell membrane, eukaryotic mitosis is usually characterized by the
presence of many linear chromosomes, whose kinetochores attaches to the microtubules of the
spindle. In relation to the forms of mitosis, closed intranuclear pleuromitosis seems to be the
most primitive type, as it is more similar to bacterial division.[11]

Gallery

Mitotic cells can be visualized microscopically by staining them with fluorescent antibodies and
dyes.
Early prophase: Polar microtubules, shown as green strands, have established a matrix around the
currently intact nucleus, with the condensing chromosomes in blue. The red nodules are the
centromeres.

Early prometaphase: The nuclear membrane has just disassembled, allowing the microtubules to
quickly interact with the kinetochores, which assemble on the centromeres of the condensing
chromosomes.
Metaphase: The centrosomes have moved to the poles of the cell and have established the mitotic
spindle. The chromosomes have congressed at the metaphase plate.

Anaphase: Kinetochore microtubules pull the two sets of chromosomes apart, and lengthening
polar microtubules push the halves of the dividing cell further apart, while chromosomes are
condensed maximally.
Telophase: Reversal of prophase and prometaphase events and thus completing the cell cycle.

See also

Aneuploidy

Binary fission

Chromosome abnormality

Cytoskeleton

Meiosis

Mitogen

Mitosis Promoting Factor

Mitotic bookmarking

Motor protein

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Further reading

Morgan DL (2007). The cell cycle: principles of control. London: Published by New Science Press in
association with Oxford University Press. ISBN 978-0-9539181-2-6.

Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "Mitosis" (https://www.ncbi.nlm.nih.go

v/books/NBK26934/) . Molecular Biology of the Cell (4th ed.). Garland Science. Retrieved 2006-01-22.

Campbell N, Reece J (December 2001). "The Cell Cycle" (https://archive.org/details/biologyc00camp) .

Biology (https://archive.org/details/biologyc00camp/page/217) (6th ed.). San Francisco: Benjamin
Cummings/Addison-Wesley. pp. 217–224 (https://archive.org/details/biologyc00camp/page/217) .
ISBN 978-0-8053-6624-2.
Cooper G (2000). "The Events of M Phase" (https://www.ncbi.nlm.nih.gov/books/NBK9958/) . The Cell:
A Molecular Approach (2nd ed.). Sinaeur Associates, Inc. Retrieved 2006-01-22.

Freeman S (2002). "Cell Division". Biological Science

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External links

Wikimedia Commons has media related to Mitosis.

Wikiversity has learning resources about Overview of Cell Biology/Mitosis

A Flash animation comparing Mitosis and Meiosis (https://www.pbs.org/wgbh/nova/miracle/

divide.html#)

Khan Academy, lecture (https://web.archive.org/web/20111228073007/http://www.khanacad

emy.org/video/phases-of-mitosis?playlist=Biology)

Studying Mitosis in Cultured Mammalian Cells (http://www.cshprotocols.org/cgi/content/full/

2007/3/pdb.prot4674)

General K-12 classroom resources for Mitosis (http://www.lessonplanet.com/directory_article

s/biology_lesson_plans/24_April_2010/361/making_mitosis_movies)

The Cell-Cycle Ontology (https://archive.today/20120730222759/http://www.semantic-syste

ms-biology.org/apo/)

WormWeb.org: Interactive Visualization of the C. elegans Cell Lineage (http://wormweb.org/ce

lllineage) – Visualize the entire cell lineage tree and all of the cell divisions of the nematode
C. elegans
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