Microtubules, like microfilaments, are dynamic structures, changing in length and interactions to react to intra- and extra-cellular changes. However, the general placement of microtubules within the cell is significantly different from microfilaments, although there is some overlap as well as interaction. Microfilaments do not have any kind of global organization with respect to their polarity. They start and end in many areas of the cell. On the other hand, almost all microtubules have their (-) end in a perinuclear area known as the MTOC, or microtubule organizing center and they radiate outward from that center. Since the microtubules all radiate outward from the MTOC, it is not surprising that they are concentrated more centrally in the cell than the microfilaments which, as mentioned above, are more abundant around the periphery of the cell. In some cell types (primarily animal), the MTOC contains a structure known as the centrosome. This consists of a centriole (two short barrel-shaped microtubule- based structures positioned perpendicular to each other) and a poorly defined concentration of pericentriolar material (PCM). The centriole is composed of nine fibrils, all connected to form a cylinder, and each also connected by radial spokes to a central axis. The electron micrograph in figure 5 shows a cross-section of a centriole. In it, each fibril is shown to actually be a fused triplet of microtubules.
Figure 5. An electron micrograph depicting the cross-section of a centriole in an embryonic mouse brain cell. L. Howard and M. Marin-Padilla, 1985
Inhibition of γ-tubulin function by antibody blocking, RNA interference of expression, and gene knockout confirm that without γ-tubulin function, the microtubule structures did not form. In addition, it appears to be play roles in coordination of late mitosis (anaphase onwards).
However, in each triplet, only one is a complete microtubule (designated the A tubule), while the B and C tubules do not form complete tubes (they share a wall with the A and B tubules, respectively). Interestingly, the centrioles do not appear to be connected to the cellular microtubule network. However, whether there is a defined centrosome or not, the MTOC region is the point of origin for all microtubule arrays. This is because the MTOC contains a high concentration of γ-tubulin. Why is this important? With all of the cytoskeletal elements, though it is most pronounced with microtubules, the rate of nucleation, or starting a microtubule is significantly slower than the rate of elongating an existing structure. Since it is the same biochemical interaction, the assumption is that the difficulty lies in getting the initial ring of dimers into position. The γ-tubulin facilitates this process by forming a γ-tubulin ring complex that serves as a template for the nucleation of microtubules (fig. 6).
Figure 6. γ-tubulin ring complex facilitates microtubule nucleation.
This is true both in animal and fungal cells with a single defined MTOC, as well as in plant cells, which have multiple, dispersed sites of microtubule nucleation.