Cytokinesis
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Cytokinesis () is the part of the
cell division Cell division is the process by which a parent cell (biology), 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 eukar ...
process A process is a series or set of activities that interact to produce a result; it may occur once-only or be recurrent or periodic. Things called a process include: Business and management *Business process, activities that produce a specific se ...
during which 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. The ...
of a single
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 ...
cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division in
mitosis In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...
and
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 ...
. During cytokinesis 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 p ...
partitions and transports duplicated
chromatid A chromatid (Greek ''khrōmat-'' 'color' + ''-id'') is one half of a duplicated chromosome. Before replication, one chromosome is composed of one DNA molecule. In replication, the DNA molecule is copied, and the two molecules are known as chro ...
s into the cytoplasm of the separating daughter cells. It thereby ensures that
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 ...
number and complement are maintained from one generation to the next and that, except in special cases, the daughter cells will be functional copies of the parent cell. After the completion of the
telophase Telophase () is the final stage in both meiosis and mitosis in a eukaryotic cell. During telophase, the effects of prophase and prometaphase (the nucleolus and nuclear membrane disintegrating) are reversed. As chromosomes reach the cell poles, ...
and cytokinesis, each daughter cell enters the
interphase Interphase is the portion of the cell cycle that is not accompanied by visible changes under the microscope, and includes the G1, S and G2 phases. During interphase, the cell grows (G1), replicates its DNA (S) and prepares for mitosis (G2). A c ...
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 ...
. Particular functions demand various deviations from the process of symmetrical cytokinesis; for example in
oogenesis Oogenesis, ovogenesis, or oögenesis is the differentiation of the ovum (egg cell) into a cell competent to further develop when fertilized. It is developed from the primary oocyte by maturation. Oogenesis is initiated in the embryonic stage. O ...
in animals the ovum takes almost all the cytoplasm and
organelles 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 ...
. This leaves very little for the resulting
polar bodies A polar body is a small haploid cell that is formed at the same time as an egg cell during oogenesis, but generally does not have the ability to be fertilized. It is named from its polar position in the egg. When certain diploid cells in animals ...
, which in most species die without function, though they do take on various special functions in other species. Another form of mitosis occurs in tissues such as
liver The liver is a major Organ (anatomy), organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of proteins and biochemicals necessary for ...
and
skeletal muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
; it omits cytokinesis, thereby yielding
multinucleate 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. Plant cytokinesis differs from animal cytokinesis, partly because of the rigidity of plant cell walls. Instead of plant cells forming a cleavage furrow such as develops between animal daughter cells, a dividing structure known as the
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 in the cytoplasm and grows into a new, doubled
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
between plant daughter cells. It divides the cell into two daughter cells. Cytokinesis largely resembles the
prokaryotic A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...
process of
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 ta ...
, but because of differences between prokaryotic and eukaryotic cell structures and functions, the mechanisms differ. For instance, a bacterial cell has a
Circular chromosome A circular chromosome is a chromosome in bacteria, archaea, Mitochondrial DNA#Genome structure and diversity, mitochondria, and Chloroplast DNA#Molecular structure, chloroplasts, in the form of a molecule of circular DNA, unlike the linear chromo ...
(a single chromosome in the form of a closed loop), in contrast to the
linear Linearity is the property of a mathematical relationship (''function'') that can be graphically represented as a straight line. Linearity is closely related to '' proportionality''. Examples in physics include rectilinear motion, the linear r ...
, usually multiple, chromosomes of eukaryote. Accordingly, bacteria construct no mitotic spindle in cell division. Also, duplication of prokaryotic DNA takes place during the actual separation of chromosomes; in mitosis, duplication takes place during the
interphase Interphase is the portion of the cell cycle that is not accompanied by visible changes under the microscope, and includes the G1, S and G2 phases. During interphase, the cell grows (G1), replicates its DNA (S) and prepares for mitosis (G2). A c ...
before mitosis begins, though the daughter
chromatid A chromatid (Greek ''khrōmat-'' 'color' + ''-id'') is one half of a duplicated chromosome. Before replication, one chromosome is composed of one DNA molecule. In replication, the DNA molecule is copied, and the two molecules are known as chro ...
s do not separate completely before the
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 ...
.


Etymology and pronunciation

The word "cytokinesis" () uses combining forms of '' cyto-'' + '' kine-'' + '' -sis'',
New Latin New Latin (also called Neo-Latin or Modern Latin) is the revival of Literary Latin used in original, scholarly, and scientific works since about 1500. Modern scholarly and technical nomenclature, such as in zoological and botanical taxonomy ...
from
Classical Latin Classical Latin is the form of Literary Latin recognized as a literary standard by writers of the late Roman Republic and early Roman Empire. It was used from 75 BC to the 3rd century AD, when it developed into Late Latin. In some later periods ...
and
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
, reflecting "
cell 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 ...
" and '' kinesis'' ("motion, movement"). It was coined by
Charles Otis Whitman Charles Otis Whitman (December 6, 1842 – December 14, 1910) was an American zoologist, who was influential to the founding of classical ethology (study of animal behavior). A dedicated educator who preferred to teach a few research students at ...
in 1887. Origin of this term is from
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 ...
(', a hollow), Latin derivative ' (cellular), Greek (', movement).


Animal cell

Animal cell cytokinesis begins shortly after the onset of sister
chromatid A chromatid (Greek ''khrōmat-'' 'color' + ''-id'') is one half of a duplicated chromosome. Before replication, one chromosome is composed of one DNA molecule. In replication, the DNA molecule is copied, and the two molecules are known as chro ...
separation in the
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 ...
of
mitosis In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...
. The process can be divided to the following distinct steps: anaphase spindle reorganization, division plane specification, actin-myosin ring assembly and contraction, and abscission. Faithful partitioning of the genome to emerging daughter cells is ensured through the tight temporal coordination of the above individual events by molecular signaling pathways.


Anaphase spindle reorganization

Animal cell cytokinesis starts with the stabilization of microtubules and reorganization of the mitotic spindle to form the central spindle. 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 bun ...
(or ''spindle midzone'') forms when non-kinetochore microtubule fibers are bundled between the spindle poles. A number of different species including '' H. sapiens'', ''
D. melanogaster ''Drosophila melanogaster'' is a species of fly (the taxonomic order Diptera) in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the " vinegar fly" or "pomace fly". Starting with ...
'' and ''
C. elegans ''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek ''caeno-'' (recent), ''rhabditis'' (r ...
'' require the central spindle in order to efficiently undergo cytokinesis, although the specific
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological proper ...
associated with its absence varies from one species to the next (for example, certain Drosophila cell types are incapable of forming a cleavage furrow without the central spindle, whereas in both ''C. elegans''
embryos 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 sper ...
and human
tissue culture Tissue culture is the growth of tissues or cells in an artificial medium separate from the parent organism. This technique is also called micropropagation. This is typically facilitated via use of a liquid, semi-solid, or solid growth medium, su ...
cells a cleavage furrow is observed to form and ingress, but then regress before cytokinesis is complete). The process of mitotic spindle reorganization and central spindle formation is caused by the decline of CDK1 activity during anaphase. The decline of CDK1 activity at the metaphase-anaphase transition leads to dephosphorylating of inhibitory sites on multiple central spindle components. First of all, the removal of a CDK1 phosphorylation from a subunit of the CPC (the chromosomal passenger complex) allows its translocalization to the central spindle from the centromeres, where it is located during metaphase. Besides being a structural component of the central spindle itself, CPC also plays a role in the phosphoregulation of other central spindle components, including PRC1 (microtubule-bundling protein required for cytokinesis 1) and MKLP1 (a kinesin motor protein). Originally inhibited by CDK1-mediated phosphorylation, PRC1 is now able to form a homodimer that selectively binds to the interface between antiparallel microtubules, facilitating spatial organization of the microtubules of the central spindle. MKLP1, together with the Rho-family GTPase activating protein CYK-4 (also termed MgcRacGAP), forms the centralspindlin complex. Centralspindlin binds to the central spindle as higher-order clusters. The centralspindlin cluster formation is promoted by phosphorylation of MLKP1 by Aurora B, a component of CPC. In short, the self-assembly of central spindle is initiated through the phosphoregulation of multiple central spindle components by the decline of CDK1 activity, either directly or indirectly, at the metaphase-anaphase transition. The central spindle may have multiple functions in cytokinesis including the control of cleavage furrow positioning, the delivery of membrane vesicles to the cleavage furrow, and the formation of the midbody structure that is required for the final steps of division.


Division plane specification

The second step of animal cell cytokinesis involves division plane specification and cytokinetic furrow formation. Precise positioning of the division plane between the two masses of segregated chromosomes is essential to prevent chromosome loss. Meanwhile, the mechanism by which the spindle determines the division plane in animal cells is perhaps the most enduring mystery in cytokinesis and a matter of intense debate. There exist three hypotheses of furrow induction. The first is the astral stimulation hypothesis, which postulates that astral microtubules from the spindle poles carry a furrow-inducing signal to the cell cortex, where signals from two poles are somehow focused into a ring at the spindle. A second possibility, called the central spindle hypothesis, is that the cleavage furrow is induced by a positive stimulus that originates in the central spindle equator. The central spindle may contribute to the specification of the division plane by promoting concentration and activation of the small GTPase RhoA at the equatorial cortex. A third hypothesis is the astral relaxation hypothesis. It postulates that active actin-myosin bundles are distributed throughout the cell cortex, and inhibition of their contraction near the spindle poles results in a gradient of contractile activity that is highest at the midpoint between poles. In other words, astral microtubules generate a negative signal that increases cortical relaxation close to the poles. Genetic and laser-micromanipulation studies in C. elegans embryos have shown that the spindle sends two redundant signals to the cell cortex, one originating from the central spindle, and a second signal deriving from the spindle aster, suggesting the involvement of multiple mechanisms combined in the positioning of the cleavage furrow. The predominance of one particular signal varies between cell types and organisms. And the multitude and partial redundancy of signals may be required to make the system robust and to increase spatial precision.


Actin-myosin ring assembly and contraction

At the cytokinesis furrow, it is the actin-myosin contractile ring that drives the cleavage process, during which cell membrane and wall grow inward, which eventually pinches the mother cell in two. The key components of this ring are the filamentous protein actin and the motor protein myosin II. The contractile ring assembles equatorially (in the middle of the cell) at the
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 ...
(adjacent to the cell membrane). Rho protein family (RhoA protein in mammalian cells) is a key regulator of contractile ring formation and contraction in animal cells. The RhoA pathway promotes assembly of the actin-myosin ring by two main effectors. First, RhoA stimulates nucleation of unbranched actin filaments by activation of Diaphanous-related formins. This local generation of new actin filaments is important for the contractile ring formation. This actin filament formation process also requires a protein called profilin, which binds to actin monomers and helps load them onto the filament end. Second, RhoA promotes myosin II activation by the kinase ROCK, which activates myosin II directly by phosphorylation of the myosin light chain and also inhibits myosin phosphatase by phosphorylation of the phosphatase-targeting subunit MYPT. Besides actin and myosin II, the contractile ring contains the scaffolding protein anillin. Anillin binds to actin, myosin, RhoA, and CYK-4, and thereby links the equatorial cortex with the signals from the central spindle. It also contributes to the linkage of the actin-myosin ring to the plasma membrane. Additionally, anillin generates contractile forces by rectifying thermal fluctuations. Another protein, septin, has also been speculated to serve as a structural scaffold on which the cytokinesis apparatus is organized. Following its assembly, contraction of the actin-myosin ring leads to ingression of the attached plasma membrane, which partitions the cytoplasm into two domains of emerging sister cells. The force for the contractile processes is generated by movements along actin by the motor protein myosin II. Myosin II uses the free energy released when ATP is hydrolyzed to move along these actin filaments, constricting the cell membrane to form 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. ...
. Continued
hydrolysis Hydrolysis (; ) is any chemical reaction in which a molecule of water breaks one or more chemical bonds. The term is used broadly for substitution reaction, substitution, elimination reaction, elimination, and solvation reactions in which water ...
causes this cleavage furrow to ingress (move inwards), a striking process that is clearly visible 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 ...
.


Abscission

The cytokinetic furrow ingresses until a midbody structure (composed of electron-dense, proteinaceous material) is formed, where the actin-myosin ring has reached a diameter of about 1–2 μm. Most animal cell types remain connected by an intercellular cytokinetic bridge for up to several hours until they are split by an actin-independent process termed abscission, the last step of cytokinesis. The process of abscission physically cleaves the midbody into two. Abscission proceeds by removal of cytoskeletal structures from the cytokinetic bridge, constriction of the cell cortex, and plasma membrane fission. The intercellular bridge is filled with dense bundles of antiparallel microtubules that derive from the central spindle. These microtubules overlap at the midbody, which is generally thought to be a targeting platform for the abscission machinery. The microtubule severing protein
spastin The human gene SPAST codes for the microtubule-severing protein of the same name, commonly known as spastin. This gene encodes a member of the AAA (ATPases associated with a variety of cellular activities) protein family. Members of this protein ...
is largely responsible for the disassembly of microtubule bundles inside the intercellular bridge. Complete cortical constriction also requires removal of the underlying cytoskeletal structures. Actin filament disassembly during late cytokinesis depends on the PKCε–14-3-3 complex, which inactivates RhoA after furrow ingression. Actin disassembly is further controlled by the GTPase Rab35 and its effector, the phosphatidylinositol-4,5-bisphosphate 5-phosphatase OCRL. Understanding the mechanism by which the plasma membrane ultimately splits requires further investigation.


Timing cytokinesis

Cytokinesis must be temporally controlled to ensure that it occurs only after sister chromatids separate during the
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 ...
portion of normal proliferative cell divisions. To achieve this, many components of the cytokinesis machinery are highly regulated to ensure that they are able to perform a particular function at only a particular stage 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 ...
. Cytokinesis happens only after APC binds with CDC20. This allows for the separation of chromosomes and myosin to work simultaneously. After cytokinesis, non-kinetochore
microtubules 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 ...
reorganize and disappear into a new cytoskeleton as the cell cycle returns to
interphase Interphase is the portion of the cell cycle that is not accompanied by visible changes under the microscope, and includes the G1, S and G2 phases. During interphase, the cell grows (G1), replicates its DNA (S) and prepares for mitosis (G2). A c ...
(see also
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 ...
).


Plant cell

Due to the presence of a
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
, cytokinesis in plant cells is significantly different from that in animal cells, Rather than forming a contractile ring, plant cells construct 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 ...
in the middle of the cell. The stages of
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 ...
formation include (1) creation 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- ...
, an array of
microtubules 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 ...
that guides and supports the formation of the
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 ...
; (2) trafficking of vesicles to the division plane and their fusion to generate a tubular-vesicular network; (3) continued fusion of membrane tubules and their transformation into membrane sheets upon the deposition of
callose Callose is a plant polysaccharide. Its production is due to the glucan synthase-like gene (GLS) in various places within a plant. It is produced to act as a temporary cell wall in response to stimuli such as stress or damage. Callose is composed ...
, followed by deposition of
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
and other
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
components; (4) recycling of excess membrane and other material from the
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 ...
; and (5) fusion with the parental
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
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- ...
is assembled from the remnants of the
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 ...
, and serves as a track for the trafficking of
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 ...
to the phragmoplast midzone. These vesicles contain lipids, proteins and carbohydrates needed for the formation of a new cell boundary. Electron tomographic studies have identified 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 ...
as the source of these vesicles, but other studies have suggested that they contain endocytosed material as well. These tubules then widen and fuse laterally with each other, eventually forming a planar, fenestrated sheet /sup>. As the
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 ...
matures, large amounts of membrane material are removed via clathrin-mediated
endocytosis Endocytosis is a cellular process in which substances are brought into the cell. The material to be internalized is surrounded by an area of cell membrane, which then buds off inside the cell to form a vesicle containing the ingested material. E ...
/sup> Eventually, the edges of the cell plate fuse with the parental plasma membrane, often in an asymmetrical fashion, thus completing cytokinesis. The remaining fenestrae contain strands of endoplasmic reticulum passing through them, and are thought to be the precursors of
plasmodesmata Plasmodesmata (singular: plasmodesma) are microscopic channels which traverse the cell walls of plant cells and some algal cells, enabling transport and communication between them. Plasmodesmata evolved independently in several lineages, and spec ...
/sup>. The construction of the new
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
begins within the lumen of the narrow tubules of the young
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 ...
. The order in which different cell wall components are deposited has been determined largely by immuno-electron microscopy. The first components to arrive are
pectins Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
,
hemicellulose A hemicellulose (also known as polyose) is one of a number of heteropolymers (matrix polysaccharides), such as arabinoxylans, present along with cellulose in almost all terrestrial plant cell walls.Scheller HV, Ulvskov Hemicelluloses.// Annu Rev ...
s, and arabinogalactan proteins carried by the secretory vesicles that fuse to form the cell plate. The next component to be added is
callose Callose is a plant polysaccharide. Its production is due to the glucan synthase-like gene (GLS) in various places within a plant. It is produced to act as a temporary cell wall in response to stimuli such as stress or damage. Callose is composed ...
, which is polymerized directly at the cell plate by callose synthases. As the cell plate continues to mature and fuses with the parental plasma membrane, the callose is slowly replaced with
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
, the primary component of a mature cell wall /sup>. The
middle lamella The middle lamella is a layer that cements together the primary cell walls of two adjoining plant cells. It is the first formed layer to be deposited at the time of cytokinesis Cytokinesis () is the part of the cell division process during ...
(a glue-like layer containing pectin) develops from the cell plate, serving to bind the cell walls of adjoining cells together.


Forces


Animal cells

Cytokinetic furrow ingression is powered by Type II
Myosin ATPase Myosin ATPase () is an enzyme with List of enzymes, systematic name ''ATP phosphohydrolase (actin-translocating)''. This enzyme catalysis, catalyses the following chemical reaction : ATP + H2O \rightleftharpoons ADP + phosphate ATP hydrolysis pr ...
. Since Myosins are recruited to the medial region, the contractile forces acting on the cortex resemble a 'purse string' constriction pulling inwards. This leads to the inward constriction. The plasma membrane by virtue of its close association with the cortex via crosslinker proteins To the constriction of the cleavage furrow, the total surface area should be increased by supplying the plasma membrane via exocytosis
.


Proteins involved in cytokinesis

CEP55 is a mitotic phosphoprotein that plays a key role in cytokinesis, the final stage of cell division.


See also

* * * * * * * *


References


Further reading

*''The Molecular Requirements for Cytokinesis'' by M. Glotzer (2005), Science 307, 1735 *"Animal Cytokinesis: from parts list to mechanism" by Eggert, U.S., Mitchison, T.J., Field, C.M. (2006), Annual Review of Cell Biology 75, 543-66 *''Campbell Biology'' (2010), 580-582
More description and nice images of cell division in plants, with a focus on fluorescence microscopy
* {{Cell cycle Cell cycle Mitosis Meiosis