Oxymonads | |
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Monocercomonoides melolanthae | |
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Order: | Oxymonadida Grassé 1952 emend. Cavalier-Smith 2003 |
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The Oxymonads (or Oxymonadida) 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 (not even anaerobic mitochondrion-like organelles like hydrogenosomes or mitosomes) in oxymonads [1] and three species have been shown to completely lack any molecular markers of mitochondria. [2]
It includes e.g. Dinenympha , Pyrsonympha , Oxymonas , [3] Streblomastix , [4] Monocercomonoides , and Blattamonas . [5]
Most Oxymonads are around 50 μm in size and have a single nucleus, associated with four flagella. Their basal bodies give rise to several long sheets of microtubules, which form an organelle called an axostyle, but different in structure from the axostyles of parabasalids. The cell may use the axostyle to swim, as the sheets slide past one another and cause it to undulate. An associated fiber called the preaxostyle separates the flagella into two pairs. A few oxymonads have multiple nuclei, flagella, and axostyles.
The free-living flagellates Trimastix and Paratrimastix are closely related to the oxymonads. [6] [7] They lack aerobic mitochondria and have four flagella separated by a preaxostyle, but unlike the oxymonads have a feeding groove. This character places the Oxymonads, Trimastix, and Paratrimastix among the Excavata, and in particular they may belong to the metamonads. Molecular phylogenetic studies indeed place Preaxostyla (oxymonads, Trimastix , and Paratrimastix ) in Metamonada. [8] [9]
Cladogram of Oxymonadida [10] |
The Percolozoa are a group of colourless, non-photosynthetic Excavata, including many that can transform between amoeboid, flagellate, and cyst stages.
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.
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 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.
The metamonads are a large group of flagellate amitochondriate microscopic eukaryotes. They include the retortamonads, diplomonads, parabasalids, oxymonads, and a range of more poorly studied taxa, most of which are free-living flagellates. All metamonads are anaerobic, and most members of the four groups listed above are symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals.
Lobosa is a taxonomic group of amoebae in the phylum Amoebozoa. Most lobosans possess broad, bluntly rounded pseudopods, although one genus in the group, the recently discovered Sapocribrum, has slender and threadlike (filose) pseudopodia. In current classification schemes, Lobosa is a subphylum, composed mainly of amoebae that have lobose pseudopods but lack cilia or flagella.
The Archamoebae are a group of protists originally thought to have evolved before the acquisition of mitochondria by eukaryotes. They include genera that are internal parasites or commensals of animals. A few species are human pathogens, causing diseases such as amoebic dysentery. The other genera of archamoebae live in freshwater habitats and are unusual among amoebae in possessing flagella. Most have a single nucleus and flagellum, but the giant amoeba Pelomyxa has many of each.
A symbiotic eukaryote that lives in the hindgut of termites, Streblomastix is a protist associated with a community of ectosymbiotic bacteria.
Trimastix is a genus of excavate protists, the sole occupant of the order Trimastigida. Trimastix are bacterivorous, free living and anaerobic. It was first observed in 1881 by William Kent. There are few known species, and the genus's role in the ecosystem is largely unknown. However, it is known that they generally live in marine environments within the tissues of decaying organisms to maintain an anoxic environment. Much interest in this group is related to its close association with other members of Preaxostyla. These organisms do not have classical mitochondria, and as such, much of the research involving these microbes is aimed at investigating the evolution of mitochondria.
Anaeromonadea, also known as Preaxostyla, is a class of excavate protists, comprising the oxymonads, Trimastix, and Paratrimastix. This group is studied as a model system for reductive evolution of mitochondria, because it includes both organisms with anaerobic mitochondrion-like organelles, and those that have completely lost their mitochondria.
Conosa is a grouping of Amoebozoa. It is subdivided into three groups: Archamoeba, Variosea and Mycetozoa.
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.
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.
Chilomastix is a genus of pyriform excavates within the family Retortamonadidae All species within this genus are flagellated, structured with three flagella pointing anteriorly and a fourth contained within the feeding groove. Chilomastix also lacks Golgi apparatus and mitochondria but does possess a single nucleus. The genus parasitizes a wide range of vertebrate hosts, but is known to be typically non-pathogenic, and is therefore classified as harmless. The life cycle of Chilomastix lacks an intermediate host or vector. Chilomastix has a resistant cyst stage responsible for transmission and a trophozoite stage, which is recognized as the feeding stage. Chilomastix mesnili is one of the more studied species in this genus due to the fact it is a human parasite. Therefore, much of the information on this genus is based on what is known about this one species.
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.
Anaeramoeba is a genus of anaerobic protists of uncertain phylogenetic position, first described in 2016.
Stygiellidae is a family of free-living marine flagellates belonging to the order Jakobida, a deep-branching lineage within the eukaryotic supergroup Discoba. They are unicellular organisms that commonly inhabit anoxic, sulfide-rich and ammonium-rich marine habitats worldwide.
Paratrimastix is a genus of free-living freshwater anaerobic excavate protists from the group Metamonada, that was segregated from the genus Trimastix in 2015. The best studied species is Paratrimastix pyriformis.
Paratrimastix pyriformis is a species of free-living (non-parasitic) anaerobic freshwater bacteriovorous flagellated protists formerly known as Trimastix pyriformis and Tetramitus pyriformis.