Astral Microtubules
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An aster is a
cellular Cellular may refer to: *Cellular automaton, a model in discrete mathematics * Cell biology, the evaluation of cells work and more * ''Cellular'' (film), a 2004 movie *Cellular frequencies, assigned to networks operating in cellular RF bands *Cell ...
structure shaped like a star, consisting of a
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 prog ...
and its associated microtubules during the early stages of mitosis in an
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
cell. Asters do not form during mitosis in
plants Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude ...
. Astral rays, composed 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 a ...
, radiate from the centrosphere and look like a cloud. Astral rays are one variant of microtubule which comes out of the centrosome; others include kinetochore microtubules and polar microtubules. During mitosis, there are five stages of cell division:
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 Prometaphase is the phase of mitosis following prophase and preceding metaphase, in eukaryote, eukaryotic Somatic (biology), somatic Cell (biology), cells. In prometaphase, the nuclear membrane breaks apart into numerous "membrane vesicles", ...
,
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, alig ...
,
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 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, ...
. During prophase, two aster-covered centrosomes migrate to opposite sides of the
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom * Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucl ...
in preparation of mitotic spindle formation. During prometaphase there is fragmentation of the nuclear envelope and formation of the mitotic spindles. During metaphase, the kinetochore microtubules extending from each centrosome connect to 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 ...
s of the chromosomes. Next, during anaphase, the kinetochore microtubules pull the sister chromatids apart into individual
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 are ...
and pull them towards the centrosomes, located at opposite ends of the cell. This allows the cell to divide properly with each daughter cell containing full replicas of chromosomes. In some cells, the orientation of the asters determines the plane of division upon which the cell will divide.


Astral microtubules

Astral microtubules are a subpopulation of microtubules, which only exist during and immediately before mitosis. They are defined as any microtubule originating from the
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 prog ...
which does not connect to a
kinetochore 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 ...
.Mitosis
Molecular Biology of the Cell, Albert et al 4th Edition. Astral microtubules develop in the actin skeleton and interact with the cell cortex to aid in spindle orientation. They are organized into radial arrays around the centrosomes. The turn-over rate of this population of microtubules is higher than any other population. The role of astral microtubules is assisted by dyneins specific to this role. These dyneins have their light chains (static portion) attached to the cell membrane, and their globular parts (dynamic portions) attached to the microtubules. The globular chains attempt to move towards the centrosome, but as they are bound to the cell membrane, this results in pulling the centrosomes towards the membrane, thus assisting
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 ...
. Astral microtubules are not required for the progression of mitosis, but they are required to ensure the fidelity of the process. The function of astral microtubules can be generally considered as determination of cell geometry. They are absolutely required for correct positioning and orientation of the mitotic
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 ...
, and are thus involved in determining the cell division site based on the geometry and polarity of the cells. The maintenance of astral microtubules is dependent on centrosomal integrity. It is also dependent on several microtubule-associated proteins such as EB1 and
adenomatous polyposis coli Adenomatous polyposis coli (APC) also known as deleted in polyposis 2.5 (DP2.5) is a protein that in humans is encoded by the ''APC'' gene. The APC protein is a negative regulator that controls beta-catenin concentrations and interacts with E-c ...
(APC). Growth of Microtubules Polymerization and nucleation are the two microscopic processes in with the growth of asters occur. At the negative ends of the aster , centrosomes will nucleate (form a nucleus) and anchor to the microtubules. At the positive end, polymerization of the aster will occur towards the outer. Cortical Dynein, a motor protein moves along the microtubules of the cell and plays a key role in the growth and inhibition of aster microtubules. A dynein that is barrier-attached can inhibit and trigger growth.


References

* Ishihara, Keisuke, et al. "Physical basis of large microtubule aster growth." ''eLife'', vol. 5, 2016. ''Gale OneFile: Health and Medicine'', link.gale.com/apps/doc/A476395269/HRCA?u=cuny_hunter&sid=HRCA&xid=5e6ad228. Accessed 28 Apr. 2021. * Laan, Liedewij et al. “Cortical Dynein Controls Microtubule Dynamics to Generate Pulling Forces That Position Microtubule Asters.” ''Cell (Cambridge)'' 148.3 (2012): 502–514. Web.


See also

* Mitosis *
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 prog ...
* Centriole *
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 ...
Cell biology Cell cycle Mitosis {{cell-cycle-stub