Metamonad

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Metamonad
Giardia lamblia.jpg
Giardia lamblia , a parasitic diplomonad
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
(unranked): Excavata
Phylum: Metamonada
Grassé 1952 emend. Cavalier-Smith 2003
Classes & orders
Synonyms [2]
  •  ?Archezoa
  • Centrosomea Chatton Villeneuve 1937
  • Metamonadina Grassé 1952
  • Polymastigota Butschli 1884
  • Tetramastigota Hulsmann & Hausmann 1994

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 (many being aerotolerant anaerobes), occurring mostly as symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals. [2]

Contents

Characteristics

A number of parabasalids and oxymonads are found in termite guts, and play an important role in breaking down the cellulose found in wood. Some other metamonads are parasites.

These flagellates are unusual in lacking mitochondria. Originally they were considered among the most primitive eukaryotes, diverging from the others before mitochondria appeared. However, they are now known to have lost mitochondria secondarily, and retain both organelles and nuclear genes derived from them. Mitochondrial relics include hydrogenosomes, which produce hydrogen, and small structures called mitosomes.

It now appears the Metamonada are, together with Malawimonas , sister clades of the Podiata. [3]

All of these groups are united by having flagella or basal bodies in characteristic groups of four, which are often associated with the nucleus, forming a structure called a karyomastigont. In addition, the genera Carpediemonas and Trimastix are now known to be close relatives of the retortamonad-diplomonad line and the oxymonads, respectively. Both are free-living and amitochondriate.

Classification

The metamonads were thought to make up part of the Excavata, a paraphyletic eukaryotic supergroup including flagellates with feeding grooves and their close relatives. Their relationships are uncertain, [4] and they do not always appear together on molecular trees. It is possible that the metamonads as defined here do not form a monophyletic subgroup.

The following higher level treatment from 2013 is based on works of Cavalier-Smith [5] with amendments within Fornicata from Yubukia, Simpson & Leander. [6]

Metamonada were once again proposed to be basal eukaryotes in 2018. [7]

Related Research Articles

<span class="mw-page-title-main">Percolozoa</span> Phylum of Excavata

The Percolozoa are a group of colourless, non-photosynthetic Excavata, including many that can transform between amoeboid, flagellate, and cyst stages.

<span class="mw-page-title-main">Stramenopile</span> Clade of eukaryotes

The Stramenopiles, also called Heterokonts, are a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have been secondarily lost. Stramenopiles represent one of the three major clades in the SAR supergroup, along with Alveolata and Rhizaria.

<span class="mw-page-title-main">Excavata</span> Supergroup of unicellular organisms belonging to the domain Eukaryota

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.

<span class="mw-page-title-main">Cercozoa</span> Group of single-celled organisms

Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead united by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eukaryotic group to be recognized mainly through molecular phylogenies. They are the natural predators of many species of bacteria. They are closely related to the phylum Retaria, comprising amoeboids that usually have complex shells, and together form a supergroup called Rhizaria.

<span class="mw-page-title-main">Parabasalid</span> Group of flagellated protists

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.

<span class="mw-page-title-main">Tectofilosid</span> Group of protists

The tectofilosids are a group of filose amoebae with shells. These are composed of organic materials and sometimes collected debris, in contrast to the euglyphids, which produce shells from siliceous scales. The shell usually has a single opening, but in Amphitrema and a few other genera it has two on opposite ends. The cell itself occupies most of the shell. They are most often found on marsh plants such as Sphagnum.

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 3 species have been shown to completely lack any molecular markers of mitochondria.

<span class="mw-page-title-main">Apusozoa</span> Phylum of micro-organisms

The Apusozoa are a paraphyletic phylum of flagellate eukaryotes. They are usually around 5–20 μm in size, and occur in soils and aquatic habitats, where they feed on bacteria. They are grouped together based on the presence of an organic shell or theca under the dorsal surface of the cell.

<span class="mw-page-title-main">Archamoebae</span> Phylum of protists

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.

Carpediemonas is genus of Metamonada, and belongs to the group Excavata. This organism is a unicellular flagellated eukaryote that was first discovered in substrate samples from the Great Barrier Reef. Carpediemonas can be found in anaerobic intertidal sediment, where it feeds on bacteria. A feature of this species is the presence of a feeding groove, a characteristic of the excavates. Like most other metamonads, Carpediemonas does not rely on an aerobic mitochondrion to produce energy. Instead, it contains hydrogenosomes that are used to produce ATP. This organism has two flagella: a posterior one used for feeding on the substrate, and an anterior one that moves in a slower sweeping motion. Carpediemonas is assigned to the fornicates, where similar Carpediemonas-like organisms are used in researching the evolution within excavates. Although Carpediemonas is a member of the metamonads, it is unusual in the sense that it is free-living and has three basal bodies.

<span class="mw-page-title-main">Trichozoa</span>

Trichozoa is a group of excavates.

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.

<span class="mw-page-title-main">Jakobid</span>

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.

<span class="mw-page-title-main">Thecofilosea</span> Class of single-celled organisms

Thecofilosea is a class of unicellular testate amoebae belonging to the phylum Cercozoa. They are amoeboflagellates, organisms with flagella and pseudopodia, distinguished from other cercozoa by their scale-lacking test composed of organic material. They are closely related to the Imbricatea, a group of testate amoebae with tests composed of inorganic silica scales.

<span class="mw-page-title-main">Sarcomonadea</span> Class of flagellate protists

The sarcomonads or class Sarcomonadea are a group of amoeboid biciliate protists in the phylum Cercozoa. They are characterized by a propensity to move through gliding on their posterior cilium or through filopodia, a lack of scales or external theca, a soft cell surface without obvious cortical filamentous or membranous skeleton, two cilia without scales or hairs, tubular mitochondrial cristae, near-spherical extrusomes, and a microbody attached to the nucleus.

Tetratrichomonas undula is a species of parabasalid.

<span class="mw-page-title-main">Varisulca</span> Proposed phylum of protists

Varisulca was a proposed basal Podiate taxon. It encompassed several lineages of heterotrophic protists, most notably the ancyromonads (planomonads), collodictyonids (diphylleids), rigifilids and mantamonadids. Recent evidence suggests that the latter three are closely related to each other, forming a clade called CRuMs, but that this is unlikely to be specifically related to ancyromonads

Paratrimastix is a genus of free-living, anaerobic, excavate protists from the group Metamonada, that was segregated from the genus Trimastix in 2015. The best studied species is Paratrimastix pyriformis.

References

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