Parabasalid | |
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Two trophozoites of Trichomonas vaginalis stained with Giemsa | |
Scientific classification | |
Domain: | Eukaryota |
Phylum: | Metamonada |
Subphylum: | Trichozoa |
(unranked): | Parabasalia Honigberg 1973 |
Orders [1] | |
The parabasalids are a group of flagellated protists within the supergroup Excavata. Most of these eukaryotic organisms form a symbiotic relationship in animals. These include a variety of forms found in the intestines of termites [2] and cockroaches, many of which have symbiotic bacteria that help them digest cellulose in woody plants. Other species within this supergroup are known parasites, and include human pathogens. [3]
The flagella are arranged in one or more clusters near the anterior of the cell. Their basal bodies are linked to parabasal fibers that attach to prominent Golgi complexes, distinctive to the group. Usually they also give rise to a sheet of cross-like microtubules that runs down the center of the cell and in some cases projects past the end. This is called the axostyle, but is different in structure from the axostyles of oxymonads.
Parabasalids are anaerobic, and lack mitochondria, but this is now known to be a result of secondary loss, and they contain small hydrogenosomes which apparently developed from reduced mitochondria. [4] Similar relics have been found in other amitochondriate flagellates, and the parabasalids are probably related to them, forming a group called the metamonads. They lack the feeding grooves found in most others, but this is probably a secondary loss as well.
Before reclassification, the parabasalids were divided into about seven [5] to 10 orders depending on sources. Present classification divides Parabasalia into four orders, that is, Trichonymphida, Spirotrichonymphida, Cristamonadida, and Trichomonadida. [1]
The parabsalid Trichomonas vaginalis is not known to undergo meiosis. However, Malik et al. [7] examined T. vaginalis for the presence of 29 genes that function in meiosis and found 27 such genes, including eight genes specific to meiosis in model organisms. These findings suggested that the capability for meiosis, and hence sexual reproduction, was likely present in a recent parabasalid ancestor of T. vaginalis. [7]
Excavata is an extensive and diverse but paraphyletic group of unicellular Eukaryota. The group was first suggested by Simpson and Patterson in 1999 and the name latinized and assigned a rank by Thomas Cavalier-Smith in 2002. It contains a variety of free-living and symbiotic protists, and includes some important parasites of humans such as Giardia and Trichomonas. Excavates were formerly considered to be included in the now obsolete Protista kingdom. They were distinguished from other lineages based on electron-microscopic information about how the cells are arranged. They are considered to be a basal flagellate lineage.
Trichomonas vaginalis is an anaerobic, flagellated protozoan parasite and the causative agent of a sexually transmitted disease called trichomoniasis. It is the most common pathogenic protozoan that infects humans in industrialized countries. Infection rates in men and women are similar but women are usually symptomatic, while infections in men are usually asymptomatic. Transmission usually occurs via direct, skin-to-skin contact with an infected individual, most often through vaginal intercourse. The WHO has estimated that 160 million cases of infection are acquired annually worldwide. The estimates for North America alone are between 5 and 8 million new infections each year, with an estimated rate of asymptomatic cases as high as 50%. Usually treatment consists of metronidazole and tinidazole.
The diplomonads are a group of flagellates, most of which are parasitic. They include Giardia duodenalis, which causes giardiasis in humans. They are placed among the metamonads, and appear to be particularly close relatives of the retortamonads.
Trichomonas is a genus of anaerobic excavate parasites of vertebrates. It was first discovered by Alfred François Donné in 1836 when he found these parasites in the pus of a patient suffering from vaginitis, an inflammation of the vagina. Donné named the genus from its morphological characteristics. The prefix tricho- originates from the Ancient Greek word θρίξ (thrix) meaning hair, describing Trichomonas’s flagella. The suffix -monas, describes its similarity to unicellular organisms from the genus Monas.
The metamonads are a large group of flagellate amitochondriate microscopic eukaryotes. Their composition is not entirely settled, but they include the retortamonads, diplomonads, and possibly the parabasalids and oxymonads as well. These four groups are all anaerobic, occurring mostly as symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals.
A hydrogenosome is a membrane-enclosed organelle found in some anaerobic ciliates, flagellates, and fungi. Hydrogenosomes are highly variable organelles that have presumably evolved from protomitochondria to produce molecular hydrogen and ATP in anaerobic conditions.
The Oxymonads are a group of flagellated protists found exclusively in the intestines of animals, mostly termites and other wood-eating insects. Along with the similar parabasalid flagellates, they harbor the symbiotic bacteria that are responsible for breaking down cellulose. There is no evidence for presence of mitochondria in oxymonads and three species have been shown to completely lack any molecular markers of mitochondria.
Trichomonadida is an order of anaerobic protists, included with the parabasalids. Members of this order are referred to as trichomonads.
An axostyle is a sheet of microtubules found in certain protists. It arises from the bases of the flagella, sometimes projecting beyond the end of the cell, and is often flexible or contractile, and so may be involved in movement and provides support for the cell. Axostyles originate in association with a flagellar microtubular root and occur in two groups, the oxymonads and parabasalids; they have different structures and are not homologous. Within trichomonads the axostyle has been theorised to participate in locomotion and cell adhesion, but also karyokinesis during cell division.
Bodo saltans is a free-living nonparasitic species of kinetoplastid flagellated phagotrophic protozoa that feed on bacteria. Bodo saltans cells have been reported in freshwater and marine environments.
Trichonympha is a genus of single-celled, anaerobic parabasalids of the order Hypermastigia that is found exclusively in the hindgut of lower termites and wood roaches. Trichonympha’s bell shape and thousands of flagella make it an easily recognizable cell. The symbiosis between lower termites/wood roaches and Trichonympha is highly beneficial to both parties: Trichonympha helps its host digest cellulose and in return receives a constant supply of food and shelter. Trichonympha also has a variety of bacterial symbionts that are involved in sugar metabolism and nitrogen fixation.
Protozoan infections are parasitic diseases caused by organisms formerly classified in the kingdom Protozoa. These organisms are now classified in the supergroups Excavata, Amoebozoa, Harosa, and Archaeplastida. They are usually contracted by either an insect vector or by contact with an infected substance or surface.
Malawimonas is genus of unicellular, heterotrophic flagellates with uncertain phylogenetic affinities. They have variably being assigned to Excavata and Loukozoa. Recent studies suggest they may be closely related to the Podiata.
Jakobids are an order of free-living, heterotrophic, flagellar eukaryotes in the supergroup Excavata. They are small, and can be found in aerobic and anaerobic environments. The order Jakobida, believed to be monophyletic, consists of only twenty species at present, and was classified as a group in 1993. There is ongoing research into the mitochondrial genomes of jakobids, which are unusually large and bacteria-like, evidence that jakobids may be important to the evolutionary history of eukaryotes.
Mastigamoeba is a genus of pelobionts, and treated by some as members of the Archamoebae group of protists. Mastigamoeba are characterized as anaerobic, amitochondriate organisms that are polymorphic. Their dominant life cycle stage is as an amoeboid flagellate. Species are typically free living, though endobiotic species have been described.
Jakoba is a genus in the taxon Excavata, and currently has a single described species, Jakoba libera described by Patterson in 1990, and named in honour of Dutch botanist Jakoba Ruinen.
Monocercomonoides is a genus of flagellate Excavata belonging to the order Oxymonadida. It was established by Bernard V. Travis and was first described as those with "polymastiginid flagellates having three anterior flagella and a trailing one originating at a single basal granule located in front of the anteriorly positioned nucleus, and a more or less well-defined axostyle". It is the first eukaryotic genus to be found to completely lack mitochondria, and all hallmark proteins responsible for mitochondrial function. The genus also lacks any other mitochondria related organelles (MROs) such as hydrogenosomes or mitosomes. Data suggests that the absence of mitochondria is not an ancestral feature, but rather due to secondary loss. Monocercomonoides sp. was found to obtain energy through an enzymatic action of nutrients absorbed from the environment. The genus has replaced the iron-sulfur cluster assembly pathway with a cytosolic sulfur mobilization system, likely acquired by horizontal gene transfer from a eubacterium of a common ancestor of oxymonads. These organisms are significant because they undermine assumptions that eukaryotes must have mitochondria to properly function. The genome of Monocercomonoides exilis has approximately 82 million base pairs, with 18 152 predicted protein-coding genes.
Stygiella /ˌstɪ.d͡ʒiˈɛ.lə/ is a genus of free-living marine flagellates belonging to the family Stygiellidae in the jakobids (excavata).
Saccinobaculus is a genus of unicellular eukaryotes that resides in the hindgut of the wood-feeding cockroach Cryptocercus punctulatus. This genus is known for its distinctive movement that resembles a snake trashing in a bag. The genus is involved in the digestion of wood materials within its insect-host and is vertically transmitted to insect progeny. The genus is the part of the family Saccinobaculidae.
Holomastigotoides is a genus of parabasalids found in the hindgut of lower termites. It is characterized by its dense, organized arrangement of flagella on the cell surface and the presence of a mitotic spindle outside its nucleus during the majority of its cell cycle. As a symbiont of termites, Holomastigotoides is able to ingest wood and aid its host in digestion. In return, Holomastigotoides is supplied with a stable habitat and steady supply of food. Holomastigotoides has notably been studied to observe the mechanisms of chromosomal pairing and segregation in haploid and diploid cells.