![Wilson1900Fig2](https://upload.wikimedia.org/wikipedia/commons/3/37/Wilson1900Fig2.jpg)
In
cell biology
Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living an ...
, mitosis () is a part of the
cell cycle
The cell cycle, or cell-division cycle, is the series of events that take place in a cell that cause it to divide into two daughter cells. These events include the duplication of its DNA (DNA replication) and some of its organelles, and sub ...
in which replicated
chromosomes
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maintained. Therefore, mitosis is also known as equational division. In general, mitosis is preceded by S phase of
interphase (during which
DNA replication
In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all living organisms acting as the most essential part for biological inheritan ...
occurs) and is often followed by
telophase and
cytokinesis
Cytokinesis () is the part of the cell division process during which the cytoplasm of a single eukaryotic cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division in mitosis and mei ...
; which divides the
cytoplasm
In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
,
organelle
In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' th ...
s and
cell membrane
The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
of one cell into two new
cells
Cell most often refers to:
* Cell (biology), the functional basic unit of life
Cell may also refer to:
Locations
* Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
containing roughly equal shares of these cellular components. 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.
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
Preprophase is an additional phase during mitosis in plant cells that does not occur in other eukaryotes such as animals or fungi. It precedes prophase and is characterized by two distinct events:
#The formation of the preprophase band, a dense ...
(specific to plant cells),
prophase
Prophase () is the first stage of cell division in both mitosis and meiosis. Beginning after interphase, DNA has already been replicated when the cell enters prophase. The main occurrences in prophase are the condensation of the chromatin ret ...
,
prometaphase,
metaphase
Metaphase ( and ) is a stage of mitosis in the eukaryotic cell cycle in which chromosomes are at their second-most condensed and coiled stage (they are at their most condensed in anaphase). These chromosomes, carrying genetic information, a ...
,
anaphase
Anaphase () is the stage of mitosis after the process of metaphase, when replicated chromosomes are split and the newly-copied chromosomes (daughter chromatids) are moved to opposite poles of the cell. Chromosomes also reach their overall maxim ...
, and
telophase. During mitosis, the chromosomes, which have already duplicated, condense and attach to spindle
fibers
Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorporate ...
that pull one copy of each chromosome to opposite sides of the cell. 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.
[ The different phases of mitosis can be visualized in real time, using live cell imaging. 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).] Other errors during mitosis can induce mitotic catastrophe
Mitotic Catastrophe has been defined as either a cellular mechanism to prevent potentially cancerous cells from proliferating or as a mode of cellular death that occurs following improper cell cycle progression or entrance. Mitotic catastrophe can ...
, apoptosis
Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes ( morphology) and death. These changes in ...
(programmed cell death) or cause mutation
In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, m ...
s. Certain types of cancer
Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal b ...
can arise from such mutations.
Mitosis occurs only in eukaryotic
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
cells. Prokaryotic cells, which lack a nucleus, divide by a different process called binary fission. Mitosis varies between organisms. For example, animal
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
cells undergo an "open" mitosis, where the nuclear envelope
The nuclear envelope, also known as the nuclear membrane, is made up of two lipid bilayer membranes that in eukaryotic cells surround the nucleus, which encloses the genetic material.
The nuclear envelope consists of two lipid bilayer membr ...
breaks down before the chromosomes separate, whereas fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
undergo a "closed" mitosis, where chromosomes divide within an intact cell nucleus.[ 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 ]gamete
A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
s – sperm
Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, ...
and egg cells – which are produced by meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
.
Discovery
Numerous descriptions of cell division
Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there ...
were made during 18th and 19th centuries, with various degrees of accuracy.[ In 1835, the German botanist ]Hugo von Mohl
Hugo von Mohl FFRS HFRSE (8 April 1805 – 1 April 1872) was a German botanist from Stuttgart. He was the first person to use the word "protoplasm".
Life
He was a son of the Württemberg statesman Benjamin Ferdinand von Mohl (1766–1845), ...
, described cell division in the green algae
The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga ...
''Cladophora glomerata
''Cladophora'' is a genus of reticulated filamentous Ulvophyceae (green algae). The genus ''Cladophora'' contains many species that are very hard to tell apart and classify, mainly because of the great variation in their appearances, which is ...
'', stating that multiplication of cells occurs through cell division. 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
Robert Remak (26 July 1815 – 29 August 1865) was a Jewish Polish-German embryologist, physiologist, and neurologist, born in Poznań, Posen, Prussia, who discovered that the origin of cells was by the Cell division, division of pre-existing cel ...
and others.
In animal cells, cell division with mitosis was discovered in frog, rabbit, and cat cornea
The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical ...
cells in 1873 and described for the first time by the Polish histologist Wacław Mayzel
Wacław Mayzel (September 12, 1847 – April 19, 1916) was a Polish histologist and the first person to describe mitosis.
Biography
Wacław Mayzel was born on September 12, 1847 to Józef and Rozalia (Minheymer) Mayzel, in Kunów, Poland. Having ...
in 1875.
Bütschli, Schneider and Fol might have also claimed the discovery of the process presently known as "mitosis".[Ross, Anna E. "Human Anatomy & Physiology I: A Chronology of the Description of Mitosis". ''Christian Brothers University''. Retrieved 02 May 2018]
link
. In 1873, the German zoologist Otto Bütschli
Johann Adam Otto Bütschli (3 May 1848 – 2 February 1920) was a German zoologist and professor at the University of Heidelberg. He specialized in invertebrates and insect development. Many of the groups of protists were first recognized by him ...
published data from observations on nematode
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s. A few years later, he discovered and described mitosis based on those observations.
The term "mitosis", coined by Walther Flemming
Walther Flemming (21 April 1843 – 4 August 1905) was a German biologist and a founder of cytogenetics.
He was born in Sachsenberg (now part of Schwerin) as the fifth child and only son of the psychiatrist Carl Friedrich Flemming (1799–18 ...
in 1882, is derived from the Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
word μίτος (''mitos'', "warp thread"). There are some alternative names for the process, e.g., "karyokinesis" (nuclear division), a term introduced by Schleicher in 1878, or "equational division", proposed by August Weismann
August Friedrich Leopold Weismann FRS (For), HonFRSE, LLD (17 January 18345 November 1914) was a German evolutionary biologist. Fellow German Ernst Mayr ranked him as the second most notable evolutionary theorist of the 19th century, after Cha ...
in 1887. However, the term "mitosis" is also used in a broad sense by some authors to refer to karyokinesis and cytokinesis together. Presently, "equational division" is more commonly used to refer to meiosis II
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately res ...
, the part of meiosis most like mitosis.
Phases
Overview
The primary result of mitosis and cytokinesis is the transfer of a parent cell's genome
In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
into two daughter cells. The genome is composed of a number of chromosomes—complexes of tightly coiled DNA that contain genetic information
A nucleic acid sequence is a succession of bases signified by a series of a set of five different letters that indicate the order of nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. By convention, sequences are u ...
vital for proper cell function. 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
S phase (Synthesis Phase) is the phase of the cell cycle in which DNA is replicated, occurring between G1 phase and G2 phase. Since accurate duplication of the genome is critical to successful cell division, the processes that occur during ...
of interphase.[ Chromosome duplication results in two identical '']sister chromatids
A sister chromatid refers to the identical copies (chromatids) formed by the DNA replication of a chromosome, with both copies joined together by a common centromere. In other words, a sister chromatid may also be said to be 'one-half' of the dup ...
'' bound together by cohesin
Cohesin is a protein complex that mediates sister chromatid cohesion, homologous recombination, and DNA looping. Cohesin is formed of SMC3, SMC1, SCC1 and SCC3 ( SA1 or SA2 in humans). Cohesin holds sister chromatids together after DNA rep ...
proteins at the ''centromere
The centromere links a pair of sister chromatids together during cell division. This constricted region of chromosome connects the sister chromatids, creating a short arm (p) and a long arm (q) on the chromatids. During mitosis, spindle fibers ...
''.
When mitosis begins, the chromosomes condense and become visible. In some eukaryotes, for example animals, the nuclear envelope
The nuclear envelope, also known as the nuclear membrane, is made up of two lipid bilayer membranes that in eukaryotic cells surround the nucleus, which encloses the genetic material.
The nuclear envelope consists of two lipid bilayer membr ...
, which segregates the DNA from the cytoplasm, disintegrates into small vesicles. The nucleolus
The nucleolus (, plural: nucleoli ) is the largest structure in the nucleus of eukaryotic cells. It is best known as the site of ribosome biogenesis, which is the synthesis of ribosomes. The nucleolus also participates in the formation of sign ...
, which makes ribosomes in the cell, also disappears. Microtubule
Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27 nm and have an inner diameter between 1 ...
s 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.[ 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 cell
Plant cells are the cells present in green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids with the capab ...
s, a cell plate
image:Phragmoplast.png, 300px, Phragmoplast and cell plate formation in a plant cell during cytokinesis. Left side: Phragmoplast forms and cell plate starts to assemble in the center of the cell. Towards the right: Phragmoplast enlarges in a donut ...
forms between the two nuclei. Cytokinesis does not always occur; coenocytic (a type of multinucleate condition) cells undergo mitosis without cytokinesis.
Interphase
The mitotic phase is a relatively short period of the cell cycle
The cell cycle, or cell-division cycle, is the series of events that take place in a cell that cause it to divide into two daughter cells. These events include the duplication of its DNA (DNA replication) and some of its organelles, and sub ...
. 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
S phase (Synthesis Phase) is the phase of the cell cycle in which DNA is replicated, occurring between G1 phase and G2 phase. Since accurate duplication of the genome is critical to successful cell division, the processes that occur during ...
. 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.[ All these phases in the cell cycle are highly regulated by ]cyclins
Cyclin is a family of proteins that controls the progression of a cell through the cell cycle by activating cyclin-dependent kinase (CDK) enzymes or group of enzymes required for synthesis of cell cycle.
Etymology
Cyclins were originally disc ...
, cyclin-dependent kinases, and other cell cycle proteins. The phases follow one another in strict order and there are "checkpoints
Checkpoint may refer to:
Places
* Border checkpoint, a place on the land border between two states where travellers and/or goods are inspected
* Security checkpoint, erected and enforced within contiguous areas under military or paramilitary cont ...
" that give the cell cues to proceed from one phase to another. 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 In population ecology, density-dependent processes occur when population growth rates are regulated by the density of a population. This article will focus on density-dependence in the context of macroparasite life cycles.
Positive density-dependen ...
) or when they differentiate to carry out specific functions for the organism, as is the case for human heart muscle cells and neurons
A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. N ...
. 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. The first process, non-homologous end joining
Non-homologous end joining (NHEJ) is a pathway that repairs double-strand breaks in DNA. NHEJ is referred to as "non-homologous" because the break ends are directly ligated without the need for a homologous template, in contrast to homology direct ...
(NHEJ), can join the two broken ends of DNA in the G1, S and G2 phases of interphase. The second process, homologous recombination
Homologous recombination is a type of genetic recombination in which genetic information is exchanged between two similar or identical molecules of double-stranded or single-stranded nucleic acids (usually DNA as in cellular organisms but may ...
al 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
In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all living organisms acting as the most essential part for biological inheritan ...
is either partially accomplished or after it is completed, since HRR requires two adjacent homologs
A couple of homologous chromosomes, or homologs, are a set of one maternal and one paternal chromosome that pair up with each other inside a cell during fertilization. Homologs have the same genes in the same locus (genetics), loci where they pr ...
.
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. Today, more research is being done to understand specifically how the phases stated above occur.
Mitosis
Preprophase
Preprophase is an additional phase during mitosis in plant cells that does not occur in other eukaryotes such as animals or fungi. It precedes prophase and is characterized by two distinct events:
#The formation of the preprophase band, a dense ...
(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
The phragmosome is a sheet of cytoplasm forming in highly vacuolated plant cells in preparation for mitosis. In contrast to animal cells, plant cells often contain large central vacuoles occupying up to 90% of the total cell volume and pushing the ...
, 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
Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of over ...
filaments (called preprophase band
The preprophase band is a microtubule array found in plant cells that are about to undergo cell division and enter the preprophase stage of the plant cell cycle. Besides the phragmosome, it is the first microscopically visible sign that a plant c ...
) underneath the plasma membrane around the equatorial plane of the future mitotic spindle
Spindle may refer to:
Textiles and manufacturing
* Spindle (textiles), a straight spike to spin fibers into yarn
* Spindle (tool), a rotating axis of a machine tool
Biology
* Common spindle and other species of shrubs and trees in genus ''Euony ...
. This band marks the position where the cell will eventually divide. The cells of higher plants (such as the flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s) lack centrioles
In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and are ...
; 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.[ The preprophase band disappears during nuclear envelope breakdown and spindle formation in prometaphase.]
Prophase
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
Chromatin is a complex of DNA and protein found in eukaryotic cells. The primary function is to package long DNA molecules into more compact, denser structures. This prevents the strands from becoming tangled and also plays important roles in r ...
. At the onset of prophase, chromatin fibers condense into discrete chromosomes that are typically visible at high magnification through a light microscope
The optical microscope, also referred to as a light microscope, is a type of microscope that commonly uses visible light and a system of lenses to generate magnified images of small objects. Optical microscopes are the oldest design of microsco ...
. 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
Transcription is the process of copying a segment of DNA into RNA. The segments of DNA transcribed into RNA molecules that can encode proteins are said to produce messenger RNA (mRNA). Other segments of DNA are copied into RNA molecules calle ...
ceases during prophase and does not resume until late anaphase to early G1 phase. The nucleolus
The nucleolus (, plural: nucleoli ) is the largest structure in the nucleus of eukaryotic cells. It is best known as the site of ribosome biogenesis, which is the synthesis of ribosomes. The nucleolus also participates in the formation of sign ...
also disappears during early prophase.
Close to the nucleus of animal cells are structures called centrosome
In cell biology, the centrosome (Latin centrum 'center' + Greek sōma 'body') (archaically cytocentre) is an organelle that serves as the main microtubule organizing center (MTOC) of the animal cell, as well as a regulator of cell-cycle progres ...
s, consisting of a pair of centriole
In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and a ...
s surrounded by a loose collection of proteins. The centrosome is the coordinating center for the cell's microtubule
Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27 nm and have an inner diameter between 11 an ...
s. 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
Tubulin in molecular biology can refer either to the tubulin protein superfamily of globular proteins, or one of the member proteins of that superfamily. α- and β-tubulins polymerize into microtubules, a major component of the eukaryotic cytoske ...
to help form a microtubule spindle apparatus. Motor proteins
Motor proteins are a class of molecular motors that can move along the cytoplasm of cells. They convert chemical energy into mechanical work by the hydrolysis of ATP. Flagellar rotation, however, is powered by a proton pump.
Cellular functions ...
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,[ and are not absolutely required for animal cell mitosis.]
Prometaphase
At the beginning of prometaphase in animal cells, phosphorylation of nuclear lamins causes the nuclear envelope
The nuclear envelope, also known as the nuclear membrane, is made up of two lipid bilayer membranes that in eukaryotic cells surround the nucleus, which encloses the genetic material.
The nuclear envelope consists of two lipid bilayer membr ...
to disintegrate into small membrane vesicles
Vesicle may refer to:
; In cellular biology or chemistry
* Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane
* Synaptic vesicle
; In human embryology
* Vesicle (embryology), bulge-like features o ...
. As this happens, microtubules invade the nuclear space. This is called ''open mitosis'', and it occurs in some multicellular organisms. Fungi and some protist
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s, such as algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
or trichomonad
Trichomonadida is an order of anaerobic protists, included with the parabasalids. Members of this order are referred to as trichomonads.
Some organisms in this order include:
*'' Trichomonas vaginalis'', an organism living inside the vagina of ...
s, undergo a variation called ''closed mitosis'' where the spindle forms inside the nucleus, or the microtubules penetrate the intact nuclear envelope.[
In late prometaphase, ''kinetochore microtubules'' begin to search for and attach to chromosomal ]kinetochores
A kinetochore (, ) is a disc-shaped protein structure associated with duplicated chromatids in eukaryotic cells where the spindle fibers attach during cell division to pull sister chromatids apart. The kinetochore assembles on the centromere and ...
.[ A ''kinetochore'' is a ]protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
aceous microtubule-binding structure that forms on the chromosomal centromere during late prophase.[ A number of ''polar microtubules'' find and interact with corresponding polar microtubules from the opposite centrosome to form the mitotic spindle.][ Although the kinetochore structure and function are not fully understood, it is known that it contains some form of ]molecular motor
Molecular motors are natural (biological) or artificial molecular machines that are the essential agents of movement in living organisms. In general terms, a motor is a device that consumes energy in one form and converts it into motion or mecha ...
.[ 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.][
]
Metaphase
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).[ 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.][ If the cell successfully passes through the metaphase checkpoint, it proceeds to anaphase.
]
Anaphase
During ''anaphase A'', the cohesin
Cohesin is a protein complex that mediates sister chromatid cohesion, homologous recombination, and DNA looping. Cohesin is formed of SMC3, SMC1, SCC1 and SCC3 ( SA1 or SA2 in humans). Cohesin holds sister chromatids together after DNA rep ...
s that bind sister chromatids together are cleaved, forming two identical daughter chromosomes. 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.[ In late anaphase, ]chromosome
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s also reach their overall maximal condensation level, to help chromosome segregation
Chromosome segregation is the process in eukaryotes by which two sister chromatids formed as a consequence of DNA replication, or paired homologous chromosomes, separate from each other and migrate to opposite poles of the nucleus. This segregation ...
and the re-formation of the nucleus. In most animal cells, anaphase A precedes anaphase B, but some vertebrate egg cells demonstrate the opposite order of events.
Telophase
Telophase (from the Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
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 is not a phase of mitosis, but rather a separate process necessary for completing cell division. In animal cells, a cleavage furrow
In cell biology, the cleavage furrow is the indentation of the cell's surface that begins the progression of cleavage, by which animal and some algal cells undergo cytokinesis, the final splitting of the membrane, in the process of cell division. ...
(pinch) containing a contractile ring, develops where the metaphase plate used to be, pinching off the separated nuclei.[ In both animal and plant cells, cell division is also driven by vesicles derived from the ]Golgi apparatus
The Golgi apparatus (), also known as the Golgi complex, Golgi body, or simply the Golgi, is an organelle found in most eukaryotic cells. Part of the endomembrane system in the cytoplasm, it packages proteins into membrane-bound vesicles ins ...
, which move along microtubules to the middle of the cell.[ In plants, this structure coalesces into a cell plate at the center of the ]phragmoplast
image:Phragmoplast.png, 300px, Phragmoplast and cell plate formation in a plant cell during cytokinesis. Left side: Phragmoplast forms and cell plate starts to assemble in the center of the cell. Towards the right: Phragmoplast enlarges in a donut- ...
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 image:Phycoplast.png, Schematic representation of types of cytokinesis in the green algae: 1) Phycoplast formation with cleavage furrow (e.g. ''Chlamydomonas''); 2) Cleavage furrow and persistent telophase spindle (e.g. ''Klebsormidium''); 3) Phycop ...
microtubule array during cytokinesis. 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
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
, slime mold
Slime mold or slime mould is an informal name given to several kinds of unrelated eukaryotic organisms with a life cycle that includes a free-living single-celled stage and the formation of spores. Spores are often produced in macroscopic mu ...
s, 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.[
]
Function
Mitosis's "function
Function or functionality may refer to:
Computing
* Function key, a type of key on computer keyboards
* Function model, a structured representation of processes in a system
* Function object or functor or functionoid, a concept of object-oriente ...
" 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
A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism.
In multicellula ...
and also the basis of the growth of a multicellular
A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism.
All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- ...
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
Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek language, Greek ''erythros'' for "red" and ''k ...
have a short lifespan (only about 3 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
Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish ...
regenerate lost arms through mitosis.
*Asexual reproduction: Some organisms produce genetically similar offspring through asexual reproduction
Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the fu ...
. For example, the hydra
Hydra generally refers to:
* Lernaean Hydra, a many-headed serpent in Greek mythology
* ''Hydra'' (genus), a genus of simple freshwater animals belonging to the phylum Cnidaria
Hydra or The Hydra may also refer to:
Astronomy
* Hydra (constel ...
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
Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is any form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or speci ...
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
The nuclear envelope, also known as the nuclear membrane, is made up of two lipid bilayer membranes that in eukaryotic cells surround the nucleus, which encloses the genetic material.
The nuclear envelope consists of two lipid bilayer membr ...
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
In cell biology, the spindle apparatus refers to the cytoskeletal structure of eukaryotic cells that forms during cell division to separate sister chromatids between daughter cells. It is referred to as the mitotic spindle during mitosis, a pr ...
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 The central spindle is a microtubule based structure, which forms in between segregating chromosomes during anaphase where the two sets of microtubules, emanating from opposite halves of the cell, overlap, and become arranged into antiparallel bundl ...
in case of closed pleuromitosis: "extranuclear" (spindle located in the cytoplasm) or "intranuclear" (in the nucleus).
File:Mitosis classification closed intranuclear pleuromitoses.svg, closed
intranuclear
pleuromitosis
File:Mitosis classification closed extranuclear pleuromitoses.svg, closed
extranuclear
pleuromitosis
File:Mitosis classification closed orthomitoses.svg, closed
orthomitosis
File:Mitosis classification semiopen pleuromitoses.svg, semiopen
pleuromitosis
File:Mitosis classification semiopen orthomitoses.svg, semiopen
orthomitosis
File:Mitosis classification open orthomitoses.svg, open
orthomitosis
Nuclear division takes place only in cells of organisms of the eukaryotic
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
domain, as bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
and archaea
Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
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
Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free- ...
, which show exclusively closed mitosis. Following, the occurrence of the forms of mitosis in eukaryotes:
*''Closed intranuclear pleuromitosis'' is typical of Foraminifera
Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
, some Prasinomonadida, some Kinetoplastida
Kinetoplastida (or Kinetoplastea, as a class) is a group of flagellated protists belonging to the phylum Euglenozoa, and characterised by the presence of an organelle with a large massed DNA called kinetoplast (hence the name). The organisms are ...
, the Oxymonadida
The Oxymonads (or Oxymonadida) are a group of flagellated protozoa found exclusively in the intestines of termites and other wood-eating insects. Along with the similar parabasalid flagellates, they harbor the symbiotic bacteria that are responsi ...
, the Haplosporidia
The Ascetosporea are a group of eukaryotes that are parasites of animals, especially marine invertebrates. The two groups, the haplosporids and paramyxids, are not particularly similar morphologically, but consistently group together on molecula ...
, many fungi (chytrid
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytrid ...
s, oomycete
Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
s, zygomycete
Zygomycota, or zygote fungi, is a former phylum, division or phylum of the kingdom Fungi. The members are now part of two Phylum, phyla: the Mucoromycota and Zoopagomycotina, Zoopagomycota. Approximately 1060 species are known. They are mostly t ...
s, ascomycete
Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defi ...
s), and some Radiolaria
The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell (biology), cell into the inner and outer portions of endoplasm and Ecto ...
(Spumellaria
Spumellaria is an order of radiolarians in the class Polycystinea. They are ameboid protists appearing in abundance in the world's oceans, possessing a radially-symmetrical silica (opal) skeleton that has ensured their preservation in fossil rec ...
and Acantharia); it seems to be the most primitive type.
*''Closed extranuclear pleuromitosis'' occurs in Trichomonadida
Trichomonadida is an order of anaerobic protists, included with the parabasalids. Members of this order are referred to as trichomonads.
Some organisms in this order include:
*''Trichomonas vaginalis'', an organism living inside the vagina of h ...
and Dinoflagellata.
* ''Closed orthomitosis'' is found among diatom
A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s, ciliate
The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...
s, some Microsporidia
Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore. They were once considered protozoans or prot ...
, unicellular yeast
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constitut ...
s and some multicellular fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
.
*''Semiopen pleuromitosis'' is typical of most Apicomplexa
The Apicomplexa (also called Apicomplexia) are a large phylum of parasitic alveolates. Most of them possess a unique form of organelle that comprises a type of non-photosynthetic plastid called an apicoplast, and an apical complex structure. T ...
.
*''Semiopen orthomitosis'' occurs with different variants in some amoebae (Lobosa
Lobosa is a taxonomic group of amoebae in the phylum Amoebozoa. Most lobosans possess broad, bluntly rounded pseudopods, although one genus in the group, the recently discovered ''Sapocribrum,'' has slender and threadlike (filose) pseudopodia. ...
) and some green flagellates (e.g., Raphidophyta or ''Volvox
''Volvox'' is a polyphyletic genus of chlorophyte green algae in the family Volvocaceae. It forms spherical colonies of up to 50,000 cells. They live in a variety of freshwater habitats, and were first reported by Antonie van Leeuwenhoek in 1700 ...
'').
*''Open orthomitosis'' is typical in mammals
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
and other Metazoa
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
, and in land plants
The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as siste ...
; but it also occurs in some protists.
Errors and other variations
Errors can occur during mitosis, especially during early embryo
An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
nic development in humans. During each step of mitosis, there are normally checkpoints as well that control the normal outcome of mitosis. But, occasionally to almost rarely, mistakes will happen. Mitotic errors can create aneuploid
Aneuploidy is the presence of an abnormal number of chromosomes in a cell, for example a human cell having 45 or 47 chromosomes instead of the usual 46. It does not include a difference of one or more complete sets of chromosomes. A cell with any ...
cells that have too few or too many of one or more chromosomes, a condition associated with cancer
Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal b ...
.[ 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.]
In ''nondisjunction
Nondisjunction is the failure of homologous chromosomes or sister chromatids to separate properly during cell division (mitosis/meiosis). There are three forms of nondisjunction: failure of a pair of homologous chromosomes to separate in meiosis I ...
'', sister chromatids fail to separate during anaphase. 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
A trisomy is a type of polysomy in which there are three instances of a particular chromosome, instead of the normal two. A trisomy is a type of aneuploidy (an abnormal number of chromosomes).
Description and causes
Most organisms that reprodu ...
'', and the latter will have only one copy, a condition known as ''monosomy
Monosomy is a form of aneuploidy with the presence of only one chromosome from a pair. Partial monosomy occurs when a portion of one chromosome in a pair is missing.
Human monosomy
Human conditions due to monosomy:
* Turner syndrome – People wit ...
''. On occasion, when cells experience nondisjunction, they fail to complete cytokinesis and retain both nuclei in one cell, resulting in binucleated cells
Binucleated cells are cells that contain two nuclei. This type of cell is most commonly found in cancer cells and may arise from a variety of causes. Binucleation can be easily visualized through staining and microscopy. In general, binucleation ...
.
''Anaphase lag Anaphase lag is a consequence of an event during cell division where sister chromatids do not properly separate from each other because of improper spindle formation. The chromosome or chromatid does not properly migrate during anaphase and the dau ...
'' occurs when the movement of one chromatid is impeded during anaphase. 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
Endoreduplication (also referred to as endoreplication or endocycling) is replication of the nuclear genome in the absence of mitosis, which leads to elevated nuclear gene content and polyploidy. Endoreplication can be understood simply as a vari ...
'' (or endoreplication) occurs when chromosomes duplicate but the cell does not subsequently divide. This results in polyploid
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...
cells or, if the chromosomes duplicates repeatedly, polytene chromosomes
Polytene chromosomes are large chromosomes which have thousands of DNA strands. They provide a high level of function in certain tissues such as salivary glands of insects.
Polytene chromosomes were first reported by E.G.Balbiani in 1881. Pol ...
. Endoreduplication is found in many species and appears to be a normal part of development
Development or developing may refer to:
Arts
*Development hell, when a project is stuck in development
*Filmmaking, development phase, including finance and budgeting
*Development (music), the process thematic material is reshaped
*Photographi ...
. 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. The cells then re-enter G1 and S phase and replicate their chromosomes again. This may occur multiple times, increasing the chromosome number with each round of replication and endomitosis. Platelet
Platelets, also called thrombocytes (from Greek θρόμβος, "clot" and κύτος, "cell"), are a component of blood whose function (along with the coagulation factors) is to react to bleeding from blood vessel injury by clumping, thereby ini ...
-producing megakaryocytes
A megakaryocyte (''mega-'' + '' karyo-'' + '' -cyte'', "large-nucleus cell") is a large bone marrow cell with a lobated nucleus responsible for the production of blood thrombocytes (platelets), which are necessary for normal blood clotting. In hum ...
go through endomitosis during cell differentiation.[
'']Amitosis
Amitosis ('' a-'' + ''mitosis''), also called karyostenosis or direct cell division or binary fission, is cell proliferation that does not occur by mitosis, the mechanism usually identified as essential for cell division in eukaryotes. The polyploi ...
'' in ciliates and in animal placental tissues results in a random distribution of parental alleles.
''Karyokinesis without cytokinesis'' originates multinucleated
Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinat ...
cells called coenocytes
A coenocyte () is a multinucleate cell (biology), cell which can result from multiple karyokinesis, nuclear divisions without their accompanying cytokinesis, in contrast to a syncytium, which results from cellular aggregation followed by dissolutio ...
.
Diagnostic marker
In histopathology
Histopathology (compound of three Greek words: ''histos'' "tissue", πάθος ''pathos'' "suffering", and -λογία '' -logia'' "study of") refers to the microscopic examination of tissue in order to study the manifestations of disease. Spe ...
, 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. 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. 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
Chromatin is a complex of DNA and protein found in eukaryotic cells. The primary function is to package long DNA molecules into more compact, denser structures. This prevents the strands from becoming tangled and also plays important roles in r ...
in the area of the mitotic figure) indicates high risk human papillomavirus infection
Human papillomavirus infection (HPV infection) is caused by a DNA virus from the ''Papillomaviridae'' family. Many HPV infections cause no symptoms and 90% resolve spontaneously within two years. In some cases, an HPV infection persists and res ...
-related Cervical cancer
Cervical cancer is a cancer arising from the cervix. It is due to the abnormal growth of cells that have the ability to invade or spread to other parts of the body. Early on, typically no symptoms are seen. Later symptoms may include abnormal ...
. In order to improve the reproducibility and accuracy of the mitotic count, automated image analysis using deep learning-based algorithms have been proposed. However, further research is needed before those algorithms can be used to routine diagnostics.
File:Normal versus atypical mitosis.jpg, Normal and atypical forms of mitosis in cancer cells. A, normal mitosis; B, chromatin bridge
Chromatin bridge is a mitotic occurrence that forms when telomeres of sister chromatids fuse together and fail to completely segregate into their respective daughter cells. Because this event is most prevalent during anaphase, the term anaphase br ...
; 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
In animal tissue, most cells round up to a near-spherical shape during mitosis. In epithelia
Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
and epidermis
The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
, an efficient rounding process is correlated with proper mitotic spindle
In cell biology, the spindle apparatus refers to the cytoskeletal structure of eukaryotic cells that forms during cell division to separate sister chromatids between daughter cells. It is referred to as the mitotic spindle during mitosis, a pr ...
alignment and subsequent correct positioning of daughter cells. 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
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
. Therefore, mitotic cell rounding is thought to play a protective role in ensuring accurate mitosis.
Rounding forces are driven by reorganization of F-actin
Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ov ...
and myosin
Myosins () are a superfamily of motor proteins best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are ATP-dependent and responsible for actin-based motility.
The first myosin ...
(actomyosin) into a contractile homogeneous cell cortex
The cell cortex, also known as the actin cortex, cortical cytoskeleton or actomyosin cortex, is a specialized layer of cytoplasmic proteins on the inner face of the cell membrane. It functions as a modulator of membrane behavior and cell surface p ...
that 1) rigidifies the cell periphery and 2) facilitates generation of intracellular hydrostatic pressure
Fluid statics or hydrostatics is the branch of fluid mechanics that studies the condition of the equilibrium of a floating body and submerged body "fluids at hydrostatic equilibrium and the pressure in a fluid, or exerted by a fluid, on an imme ...
(up to 10 fold higher than interphase). 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
In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide stru ...
. Generation of pressure is dependent on formin
Formins (formin homology proteins) are a group of proteins that are involved in the polymerization of actin and associate with the fast-growing end (barbed end) of actin filaments. Most formins are Rho-GTPase effector proteins. Formins regu ...
-mediated F-actin
Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ov ...
nucleation and Rho kinase
Rho-associated protein kinase (ROCK) is a kinase belonging to the AGC (PKA/ PKG/PKC) family of serine-threonine specific protein kinases. It is involved mainly in regulating the shape and movement of cells by acting on the cytoskeleton.
ROCKs ( ...
(ROCK)-mediated myosin II
Myosins () are a superfamily of motor proteins best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are ATP-dependent and responsible for actin-based motility.
The first myosin (M2 ...
contraction, both of which are governed upstream by signaling pathways RhoA
Transforming protein RhoA, also known as Ras homolog family member A (RhoA), is a small GTPase protein in the Rho family of GTPases that in humans is encoded by the ''RHOA'' gene. While the effects of RhoA activity are not all well known, it is ...
and ECT2
Protein ECT2 is a protein that in humans is encoded by the ''ECT2'' gene.
Function
The protein encoded by this gene is a transforming protein that is related to Rho-specific exchange factors and yeast cell cycle regulators. The expression of t ...
through the activity of Cdk1
Cyclin-dependent kinase 1 also known as CDK1 or cell division cycle protein 2 homolog is a highly conserved protein that functions as a serine/threonine protein kinase, and is a key player in cell cycle regulation. It has been highly studied in th ...
. 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-ray
An X-ray, or, much less commonly, X-radiation, is a penetrating form of high-energy electromagnetic radiation. Most X-rays have a wavelength ranging from 10 picometers to 10 nanometers, corresponding to frequencies in the range 30&nb ...
s in the G1 phase
The G1 phase, gap 1 phase, or growth 1 phase, is the first of four phases of the cell cycle that takes place in eukaryotic cell division. In this part of interphase, the cell synthesizes mRNA and proteins in preparation for subsequent steps leadi ...
of the cell cycle
The cell cycle, or cell-division cycle, is the series of events that take place in a cell that cause it to divide into two daughter cells. These events include the duplication of its DNA (DNA replication) and some of its organelles, and sub ...
repair recombinogenic DNA damages primarily by recombination between homologous chromosome
A couple of homologous chromosomes, or homologs, are a set of one maternal and one paternal chromosome that pair up with each other inside a cell during fertilization. Homologs have the same genes in the same loci where they provide points alon ...
s. Mitotic cells irradiated in the G2 phase
G2 phase, Gap 2 phase, or Growth 2 phase, is the third subphase of interphase in the cell cycle directly preceding mitosis. It follows the successful completion of S phase, during which the cell’s DNA is replicated. G2 phase ends with the o ...
repair such damages preferentially by sister-chromatid recombination. Mutation
In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...
s in gene
In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s encoding enzymes employed in recombination cause cells to have increased sensitivity to being killed by a variety of DNA damaging agents. These findings suggest that mitotic recombination is an adaptation for repairing DNA damages including those that are potentially lethal.
Evolution
There are prokaryotic homologs
A couple of homologous chromosomes, or homologs, are a set of one maternal and one paternal chromosome that pair up with each other inside a cell during fertilization. Homologs have the same genes in the same locus (genetics), loci where they pr ...
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 (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
, meiosis may have arisen after mitosis. However, sexual reproduction involving meiosis is also a primitive characteristic of eukaryotes.[Bernstein, H., Bernstein, C. Evolutionary origin and adaptive function of meiosis. In “Meiosis”, Intech Publ (Carol Bernstein and Harris Bernstein editors), Chapter 3: 41-75 (2013).] 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.
Gallery
Mitotic cells can be visualized microscopically by staining
Staining is a technique used to enhance contrast in samples, generally at the microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissues), in cytology (microscopic study of cells), and in the ...
them with fluorescent
Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation. It is a form of luminescence. In most cases, the emitted light has a longer wavelength, and therefore a lower photon energy, tha ...
antibodies
An antibody (Ab), also known as an immunoglobulin (Ig), is a large, Y-shaped protein used by the immune system to identify and neutralize foreign objects such as pathogenic bacteria and viruses. The antibody recognizes a unique molecule of the ...
and dyes
A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution and ...
.
See also
* Aneuploidy
Aneuploidy is the presence of an abnormal number of chromosomes in a cell, for example a human cell having 45 or 47 chromosomes instead of the usual 46. It does not include a difference of one or more complete sets of chromosomes. A cell with any ...
* Binary fission
Binary may refer to:
Science and technology Mathematics
* Binary number, a representation of numbers using only two digits (0 and 1)
* Binary function, a function that takes two arguments
* Binary operation, a mathematical operation that t ...
* Chromosome abnormality
A chromosomal abnormality, chromosomal anomaly, chromosomal aberration, chromosomal mutation, or chromosomal disorder, is a missing, extra, or irregular portion of chromosomal DNA. These can occur in the form of numerical abnormalities, where ther ...
* Cytoskeleton
The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compos ...
* Meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
* Mitogen
A mitogen is a small bioactive protein or peptide that induces a cell to begin cell division, or enhances the rate of division (mitosis). Mitogenesis is the induction (triggering) of mitosis, typically via a mitogen. The mechanism of action of a ...
* Mitosis Promoting Factor
Maturation-promoting factor (abbreviated MPF, also called mitosis-promoting factor or M-Phase-promoting factor) is the cyclin-Cdk complex that was discovered first in frog eggs. It stimulates the mitotic and meiotic phases of the cell cycle. MPF ...
* Mitotic bookmarking
Bookmarking (also "gene bookmarking" or "mitotic bookmarking") refers to a potential mechanism of transmission of gene expression programs through cell division.
During mitosis, gene transcription is silenced and most transcription factors are r ...
*Mitotic catastrophe
Mitotic Catastrophe has been defined as either a cellular mechanism to prevent potentially cancerous cells from proliferating or as a mode of cellular death that occurs following improper cell cycle progression or entrance. Mitotic catastrophe can ...
* Motor protein
Motor proteins are a class of molecular motors that can move along the cytoplasm of cells. They convert chemical energy into mechanical work by the hydrolysis of ATP. Flagellar rotation, however, is powered by a proton pump.
Cellular functions ...
References
Further reading
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External links
A Flash animation comparing Mitosis and Meiosis
Khan Academy, lecture
Studying Mitosis in Cultured Mammalian Cells
General K-12 classroom resources for Mitosis
The Cell-Cycle Ontology
WormWeb.org: Interactive Visualization of the ''C. elegans'' Cell Lineage
– Visualize the entire cell lineage tree and all of the cell divisions of the nematode ''C. elegans''
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Cell cycle
Articles containing video clips
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