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 progression. The centrosome provides structure for the cell. The centrosome is thought to have evolved only in the metazoan lineage of eukaryotic cells. Fungi and plants lack centrosomes and therefore use other structures to organize their microtubules. Although the centrosome has a key role in efficient mitosis in animal cells, it is not essential in certain fly and flatworm species.
Centrosomes are composed of two centrioles arranged at right angles to each other, and surrounded by a dense, highly structured mass of protein termed the pericentriolar material (PCM). The PCM contains proteins responsible for microtubule nucleation and anchoring — including γ-tubulin, pericentrin and ninein. In general, each centriole of the centrosome is based on a nine-triplet microtubule assembled in a cartwheel structure, and contains centrin, cenexin and tektin. In many cell types, the centrosome is replaced by a cilium during cellular differentiation. However, once the cell starts to divide, the cilium is replaced again by the centrosome.
History
The centrosome was discovered jointly by Walther Flemming in 1875 and Edouard Van Beneden in 1876, and later described and named in 1888 by Theodor Boveri.
Functions
Centrosomes are associated with the nuclear membrane during the prophase stage of the cell cycle. During mitosis, the nuclear membrane breaks down, and the centrosome-nucleated microtubules can interact with the chromosomes to build the mitotic spindle.
The mother centriole, the older of the two in the centriole pair, also has a central role in making cilia and flagella.
The centrosome is copied only once per cell cycle, so that each daughter cell inherits one centrosome, containing two structures called centrioles. The centrosome replicates during the S phase of the cell cycle. During the prophase in the process of cell division called mitosis, the centrosomes migrate to opposite poles of the cell. The mitotic spindle then forms between the two centrosomes. Upon division, each daughter cell receives one centrosome. Aberrant numbers of centrosomes in a cell have been associated with cancer. Doubling of a centrosome is similar to DNA replication in two respects: the semiconservative nature of the process and the action of CDK2 as a regulator of the process. But the processes are essentially different in that centrosome doubling does not occur by template reading and assembly. The mother centriole just aids in the accumulation of materials required for the assembly of the daughter centriole.
Centrioles, however, are not required for the progression of mitosis. When the centrioles are irradiated by a laser, mitosis proceeds normally with a morphologically normal spindle. Moreover, development of the fruit fly Drosophila is largely normal when centrioles are absent due to a mutation in a gene required for their duplication. In the absence of the centrioles, the microtubules of the spindle are focused by motors, allowing the formation of a bipolar spindle. Many cells can completely undergo interphase without centrioles.
Unlike centrioles, centrosomes are required for survival of the organism. Cells without centrosomes lack radial arrays of astral microtubules. They are also defective in spindle positioning and in the ability to establish a central localization site in cytokinesis. The function of centrosomes in this context is hypothesized to ensure the fidelity of cell division, because it greatly increases the efficacy. Some cell types arrest in the following cell cycle when centrosomes are absent. This is not a universal phenomenon.
When the nematode C. elegans egg is fertilized, the sperm delivers a pair of centrioles. These centrioles will form the centrosomes, which will direct the first cell division of the zygote, and this will determine its polarity. It's not yet clear whether the role of the centrosome in polarity determination is microtubule-dependent or independent.
In human reproduction, the sperm supplies the centriole that creates the centrosome and microtubule system of the zygote.
Centrosome alterations in cancer cells
Theodor Boveri, in 1914, described centrosome aberrations in cancer cells. This initial observation was subsequently extended to many types of human tumors. Centrosome alterations in cancer can be divided in two subgroups — i.e., structural or numeric aberrations — yet both can be found simultaneously in a tumor.