Ultrastructure of Centrioles and Centrosome Organization
The structural foundation of the centrosome is the centriole, which is a hollow cylinder. The wall of this cylinder is formed by nine triplets of microtubules arranged circumferentially and interconnected by protein links. Because the central part of the cylinder lacks microtubules (it is empty), the structural formula of a centriole is written as $(9 \times 3) + 0$.
A complete centrosome consists of two basic components:
- Diplosome — a pair of centrioles. In a non-dividing cell, only one diplosome is present, and its two centrioles are always oriented at right angles to each other.
- Centrosphere — a zone of clear cytoplasm immediately surrounding the diplosome.
Specialized structures known as satellites are localized within the centrosphere region. Some of these are closely apposed to the centrioles, while others lie freely in the cytoplasm. The primary function of satellites is that numerous microtubules of the cellular cytoskeleton radiate from them (rather than directly from the centriolar surface).
Reproduction (Duplication) of Centrioles
Duplication of centrosome components occurs strictly during the preparation of the cell for division. During this process, the pre-existing (mother) centriole acts as a structural template.
New organelles are formed through the polymerization of the tubulin protein and are initiated at a strict right angle to the mother centriole. The result of duplication is the appearance of two complete diplosomes (two pairs) within the cell. Notably, each diplosome always contains one parental centriole and one newly synthesized (daughter) centriole.
Axoneme: The Foundation of Flagella and Cilia
Organelles of movement, such as the cilia of the respiratory tract epithelium and the flagella of spermatozoa, are built upon an axoneme ("axial filament"). The axoneme is a derivative of cytoskeletal microtubules, but its ultrastructure differs from that of a centriole and is described by the formula $(9 \times 2) + 2$:
- Periphery: 9 doublets (pairs) of microtubules.
- Center: 1 pair of microtubules enclosed in a central sheath.
Axonemal elements are rigidly interconnected. Dynein arms (composed of the protein dynein) extend from each peripheral doublet toward the neighboring one. Additionally, radial spokes extend from the peripheral doublets to the central sheath. Externally, the axoneme is covered by the plasma membrane (plasmalemma), and the space between microtubules is filled with hyaloplasm. The overall structure of a flagellum is more complex than that of a cilium due to additional supportive structures surrounding the axial filament.
The mechanism of movement of these organelles is based on the cyclic attachment and detachment of dynein arms. This process causes adjacent microtubule doublets to slide past one another, ultimately resulting in the bending of the entire structure and the beating of the cilium or flagellum.
Basal Body
To securely anchor the axoneme to the cell cytoplasm, a basal body (kinetosome) is located in the superficial cytoplasm at the base of each cilium or flagellum.
In its ultrastructure, the basal body is completely identical to a centriole: it consists of 9 peripheral microtubule triplets (formula $(9 \times 3) + 0$) and also contains satellite particles. The transition from the basal body to the axoneme occurs in a specific manner: only two of the three microtubules from each basal body triplet extend upward to form the peripheral doublets of the axial filament.