A cytostome (from ''cyto-'', cell and ''stome-'', mouth) or cell mouth is a part of a cell specialized for

phagocytosis Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...

, usually in the form of a microtubule-supported funnel or groove. Food is directed into the cytostome, and sealed into

vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic m ...

s. Only certain groups of protozoa, such as the

Ciliophora The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a different ...

and

Excavata Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free- ...

, have cytostomes. An example is '' Balantidium coli'', a ciliate. In other protozoa, and in cells from multicellular organisms, phagocytosis takes place at any point on the cell or feeding takes place by absorption.

Structure

The cytostome forms an invagination on the cell surface and is typically directed towards the nucleus of the cell.Okuda, Kendi, et al. "The cytostome of Trypanosoma cruzi epimastigotes is associated with the flagellar complex." Experimental parasitology 92.4 (1999): 223-231. The cytostome is often labeled as the entire invagination, but in fact the cytostome only constitutes the opening of the invagination at the surface of the cell. The rest of the invagination is classified as the cytopharynx.Preston, T. M. "The Form and Function of the Cytostome-Cytopharynx of the Culture Forms of the Elasmobranch Haemoflagellate Trypanosoma raiae Laveran & Mesnil." The Journal of protozoology 16.2 (1969): 320-333. The cytopharynx works in conjunction with the cytostome in order to import macromolecules into the cell. This strong association between the cytostome and cytopharynx is often called the cytostome-cytopharynx complex or the cytopharyngeal apparatus. However, in a small number of cases the cytostome works independently in order to import macromolecules. In these instances, the cytostome imports macromolecules by directly forming vesicles that are imported into the interior of the cell. The cytostome is associated with microtubules that function in maintaining its form. One set of microtubules is arranged in a triplet formation and is located directly beneath the cytostome

membrane A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. ...

. A second set of microtubules is positioned directly beneath the flagellar pocket membrane and forms a quartet.Alcantara, Carolina L., et al. "The three-dimensional structure of the cytostome-cytopharynx complex of Trypanosoma cruzi epimastigotes." Journal of cell science 127.10 (2014): 2227-2237

Cytopharynx

Equally as important in the function of endocytosis is the structure known as the cytopharynx. The cytopharynx is a long, tube-like structure that forms the invagination associated with the cytostome. While the shape of the cytopharynx is not constant, it is typically directed towards the posterior of the cell, often hooking around a central nucleus. The length of the cytopharynx varies during the

cell cycle The cell cycle, or cell-division cycle, is the series of events that take place in a cell that cause it to divide into two daughter cells. These events include the duplication of its DNA (DNA replication) and some of its organelles, and sub ...

, however the average length is 8 μm. Much like the cytostome, a set of microtubules form an association with the cytopharynx. Two sets of microtubules follow the path of the cytopharynx in cells. These sets of microtubules form a gutter like structure that surrounds the cytopharynx. One side of the cytopharynx is not associated with these microtubules and is deemed the "nude". However, vesicles associate with this side of the cytopharynx.

Location

The location of the cytostome in most flagellated protozoa is strongly conserved. The cytostome is located on the anterior end of the cell close to a structure known as the flagellar pocket. The flagellar pocket is also an invagination in the cell and also serves as a site of endocytosis. The opening of the cytostome is approximately level with the opening of the flagellar pocket.

is a phylum of protozoa. The cytostome in this phyla can be either apical or lateral.

Function

The cytostome-cytopharynx complex functions as follows: macromolecules to be taken up by a cell enter the cytostome. Macromolecules then pass into the lumen of the cytopharynx and are transported to the posterior end of the cell where they are put into budding vesicles that are transported to others parts of the cell. The cytopharynx in this way acts much like a straw that sucks macromolecules to the posterior end of the cell. The passage of macromolecules from the entrance of the cytostome to the posterior end of the cytopharynx takes at least 2 minutes. The cytostome is the main site of endocytosis in ''Trypanosoma cruzi epimastigotes''.Porto-Carreiro, Isabel, et al. "''Trypanosoma cruzi epimastigote'' endocytic pathway: cargo enters the cytostome and passes through an early endosomal network before storage in reservosomes." European journal of cell biology 79.11 (2000): 858-869.

Associations

The cytostome has also been found to associate with the flagellum of ''

Trypanosoma cruzi ''Trypanosoma cruzi'' is a species of parasitic euglenoids. Among the protozoa, the trypanosomes characteristically bore tissue in another organism and feed on blood (primarily) and also lymph. This behaviour causes disease or the likelihood o ...

''. So far, this is the only known example of an endocytotic organelle being associated with an organelle that is used for locomotion.

Related structures

As mentioned above, the flagellar pocket is another site of endocytosis in flagellated protozoa. The flagellar pocket is an invagination that is formed around the extracellular flagellum. The flagellar pocket is a site of both endocytosis and exocytosis in cells.Eger, Iriane, and Maurilio José Soares. "Endocytosis in Trypanosoma cruzi (Euglenozoa: Kinetoplastea) epimastigotes: Visualization of ingested transferrin–gold nanoparticle complexes by confocal laser microscopy." Journal of microbiological methods 91.1 (2012): 101-105.

Visualization methods

Many methods have been used in order to visualize the cytostome. Eger et al. used gold labeled

transferrin Transferrins are glycoproteins found in vertebrates which bind to and consequently mediate the transport of iron (Fe) through blood plasma. They are produced in the liver and contain binding sites for two Fe3+ ions. Human transferrin is encode ...

molecules in combination with

confocal microscopy Confocal microscopy, most frequently confocal laser scanning microscopy (CLSM) or laser confocal scanning microscopy (LCSM), is an optical imaging technique for increasing optical resolution and contrast of a micrograph by means of using a s ...

in order to visualize the cytostome. This experiment showed that labeling with the gold particles was evident at two locations in the cells; one of the locations was the bottom of the cytopharynx, and the other location was in reservosomes in the cell. Another team used ion beam scanning electron microscopy, also known as FIB-SEM followed by three dimensional reconstruction in order to create a 3-dimensional model of the cytostome-cytopharynx complex.

References

{{Reflist Organelles