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The centrosome is the main microtubule organizing center (MTOC) of the cell as well as a regulator of cell-cycle progression. It was discovered in 1888 by Theodor Boveri and was described as the 'special organ of cell division.' Although the centrosome has a key role in efficient cell division, it has been recently shown that it is not necessary [1].

Centrosomes are composed of two orthogonally arranged centrioles surrounded by an amorphous mass of pericentriolar material (PCM). The PCM contains proteins responsible for microtubule nucleation and anchoring including γ-tubulin, pericentrin and ninein. Each centriole is comprised of nine triplet microtubule blades in a pinwheel structure as well as centrin, cenexin and tektin [1].

Higher eukaryotic cells possess a centrosome. Yeast cells have a spindle pole body (SPB) which is equivalent to metazoan centrosomes. The spindle pole body differs from the centrosome in many ways, the major difference is the lack of centrioles. Typical angiosperm plant cells do not have centrosomes or anything analogous to them in size, function or organization, but have a number of noncentrosomal MTOCs that lack centrioles. One plant is used as a model organisim for the study of the centrosome - the algae chlamydomonas.

Contents 1 Roles of the centrosome 2 Centrosome Genome 3 References 4 Additional images

Centrosomes are often associated with the nuclear membrane during interphase of the cell cycle. In 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 one that was inherited from the mother cell, also has a central role in making cilia and flagella [1].

The centrosome is duplicated only once per cell cycle so that each daughter cell inherits one centrosome, containing two centrioles. The centrosome replicates during the S phase of the cell cycle. During the prophase of mitosis, the centrosomes migrate to opposite poles of the cell. Mitotic spindle 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.

Interestingly, centrosomes are not required for the progression of mitosis. When the centrosomes 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 [4]. In the absence of the centrosome the microtubules of the spindle are focused by motors allowing the formation of a bipolar spindle. Many cells can completely undergo interphase without centrosomes [1].

Figure 1: Role of the centrosome in cell cycle progression. [2]

Although centrosomes are not required for mitosis or survival of the cell, they are required for survival of the organism. Acentrosomal cells lack radial arrays of astral microtubules. They are also defective in spindle positioning and in ability to establish a central localization site in cytokinesis. The function of centrosome in this context is hypothesized to ensure the fidelity of cell division as it is not necessary but 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 is not yet clear whether the role of the centrosome in polarity determination is microtubule dependent or independent.

Recent research in 2006 [3] indicates that centrosomes may have their own genome, previously known only in nuclei, mitochondria and chloroplasts. Unlike the latter, it is RNA-based rather than DNA-based, and apparently includes an RNA sequence capable of duplicating the centrosome genome. The putative centrosome genome RNA sequences were purified from surf clam eggs, were found in "few to no" other places in the cell, and do not appear in existing genome databases.

The existence of nucleotides associated with the centrosome remains controversial. Many studies have investigated whether nucleotides associate with the centrosome with varying results.

[1] Rieder, CL, S Faruki and A Khodjakov (2001) TRENDS in Cell Biology 11 10:413-418.

[2] Doxsey, S., W. Zimmerman and K. Mikule (2005) TRENDS in Cell Biology 15 6: 303-310.

[3] Collaboration at the Marine Biological Laboratory in Woods Hole, Mass., in June 5 2006 Proceedings of the National Academy of Sciences USA, as reported in Scientific American, p32, August 2006.

[4] Basto, R, et al. (2006) Cell 125: 1375 -186.

Proteins in different cellular compartments and structures tagged with green fluorescent protein.