Armophorea | |
---|---|
Sicuophora multigranularis (Clevelandellida) | |
Scientific classification | |
Domain: | Eukaryota |
Clade: | Diaphoretickes |
Clade: | SAR |
Clade: | Alveolata |
Phylum: | Ciliophora |
Subphylum: | Intramacronucleata |
Class: | Armophorea Lynn, 2004 |
Orders | |
Armophorea is a class of ciliates in the subphylum Intramacronucleata. [1] [2] . [3] It was first resolved in 2004 [4] and comprises three orders: Metopida, Clevelandellida, [5] and Armophorida. [6] Previously members of this class were thought to be heterotrichs because of similarities in morphology, most notably a characteristic dense arrangement of cilia surrounding their oral structures. [7] However, the development of genetic tools and subsequent incorporation of DNA sequence information has led to major revisions in the evolutionary relationships of many protists, including ciliates. [8] Metopids, clevelandellids, and armophorids were grouped into this class based on similarities in their small subunit rRNA sequences, making them one of two so-called "riboclasses" of ciliates, however, recent analyses suggest that Armophorida may not be related to the other two orders. [9]
The name Armophorea is thought to be derived from the Latin word arma, meaning weapons, or armus, meaning shoulder. This name refers to the caenomorphid members of this class, which have a characteristic military helmet-like morphology, and also a twisted appearance that looks like a shoulder. [7]
Free-living armophoreans live in anoxic or microaerobic habitats, in the sediment or water column where there is reduced or absent oxygen. Thus their distribution is quite limited, although they are found globally in both marine and freshwater habitats, as well as in terrestrial sediment. Clevelandellids live as commensal symbionts inside of the digestive tracts of terrestrial and aquatic animals. [7]
Armophoreans can survive by encystment when in unfavorable environmental conditions. This is quite important for clevelandellids because it facilitates their transmission between hosts. [7]
Like most anaerobic ciliates, armophoreans have mitochondria-derived organelles called hydrogenosomes. These specialized organelles produce energy for the cell in absence of oxygen by the fermentation of pyruvate into acetate and hydrogen. Armophoreans harbor methanogenic endosymbiotic archaea that have been located in the cytoplasm adjacent to their hydrogenosomes. Host cells can contain up to 10,000 methanogens, and they are thought to play roles in host growth and metabolism. [10] It is hypothesized that these symbionts consume the hydrogen that is produced as an end-product of fermentation, making it a more favorable reaction and increasing its energetic yield. [11] Marine armophoreans also harbor ectosymbiotic bacteria that are sulfate reducers, which are thought to play a similar role as the endosymbiotic methanogens in that they also consume hydrogen as a metabolite of host fermentation. [10]
Methanogenesis by these endosymbiotic archaea can contribute substantially to the production of methane in sulfate-rich, anoxic detrital sediments and the anoxic water column, but the contribution is modest (>2%) in sandy sediments where the ciliates are lower in number. [12] This is in contrast to clevelandellids: over 80% of the methane produced by the American cockroach can be attributed to these ciliates via their methanogenic endosymbionts. [7]
An anaerobic organism or anaerobe is any organism that does not require molecular oxygen for growth. It may react negatively or even die if free oxygen is present. In contrast, an aerobic organism (aerobe) is an organism that requires an oxygenated environment. Anaerobes may be unicellular or multicellular. Most fungi are obligate aerobes, requiring oxygen to survive. However, some species, such as the Chytridiomycota that reside in the rumen of cattle, are obligate anaerobes; for these species, anaerobic respiration is used because oxygen will disrupt their metabolism or kill them. The sea floor is possibly one of the largest accumulation of anaerobic organisms on Earth, where microbes are primarily concentrated around hydrothermal vents. These microbes produce energy in absence of sunlight or oxygen through a process called chemosynthesis, whereby inorganic compounds such as hydrogen gas, hydrogen sulfide or ferrous ions are converted into organic matter.
The Litostomatea are a class of ciliates. The group consists of three subclasses: Haptoria, Trichostomatia and Rhynchostomatia. Haptoria includes mostly carnivorous forms such as Didinium, a species of which preys primarily on the ciliate Paramecium. Trichostomatia (trichostomes) are mostly endosymbionts in the digestive tracts of vertebrates. These include the species Balantidium coli, which is the only ciliate parasitic in humans. The group Rhynchostomatia includes two free-living orders previously included among the Haptoria, but now known to be genetically distinct from them, the Dileptida and the Tracheliida.
A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and eukaryotic organisms. Most prokaryotes are unicellular and are classified into bacteria and archaea. Many eukaryotes are multicellular, but some are unicellular such as protozoa, unicellular algae, and unicellular fungi. Unicellular organisms are thought to be the oldest form of life, with early protocells possibly emerging 3.5–4.1 billion years ago.
The hypotrichs are a group of ciliated protozoa, common in fresh water, salt water, soil and moss. Hypotrichs possess compound ciliary organelles called "cirri," which are made up of thick tufts of cilia, sparsely distributed on the ventral surface of the cell. The multiple fused cilia which form a cirrus function together as a unit, enabling the organism to crawl along solid substrates such as submerged debris or sediments. Hypotrichs typically possess a large oral aperture, bordered on one side by a wreath or collar of membranelles, forming an "adoral zone of membranelles," or AZM.
Methanogens are anaerobic archaea that produce methane as a byproduct of their energy metabolism, i.e., catabolism. Methane production, or methanogenesis, is the only biochemical pathway for ATP generation in methanogens. All known methanogens belong exclusively to the domain Archaea, although some bacteria, plants, and animal cells are also known to produce methane. However, the biochemical pathway for methane production in these organisms differs from that in methanogens and does not contribute to ATP formation. Methanogens belong to various phyla within the domain Archaea. Previous studies placed all known methanogens into the superphylum Euryarchaeota. However, recent phylogenomic data have led to their reclassification into several different phyla. Methanogens are common in various anoxic environments, such as marine and freshwater sediments, wetlands, the digestive tracts of animals, wastewater treatment plants, rice paddy soil, and landfills. While some methanogens are extremophiles, such as Methanopyrus kandleri, which grows between 84 and 110°C, or Methanonatronarchaeum thermophilum, which grows at a pH range of 8.2 to 10.2 and a Na+ concentration of 3 to 4.8 M, most of the isolates are mesophilic and grow around neutral pH.
Methanogenesis or biomethanation is the formation of methane coupled to energy conservation by microbes known as methanogens. It is the fourth and final stage of anaerobic digestion. Organisms capable of producing methane for energy conservation have been identified only from the domain Archaea, a group phylogenetically distinct from both eukaryotes and bacteria, although many live in close association with anaerobic bacteria. The production of methane is an important and widespread form of microbial metabolism. In anoxic environments, it is the final step in the decomposition of biomass. Methanogenesis is responsible for significant amounts of natural gas accumulations, the remainder being thermogenic.
The hydrogen hypothesis is a model proposed by William F. Martin and Miklós Müller in 1998 that describes a possible way in which the mitochondrion arose as an endosymbiont within a prokaryotic host in the archaea, giving rise to a symbiotic association of two cells from which the first eukaryotic cell could have arisen (symbiogenesis).
Stephanopogon is a genus of flagellated marine protist that superficially resembles a ciliate.
Methanobacterium is a genus of the Methanobacteria class in the Archaea kingdom, which produce methane as a metabolic byproduct. Despite the name, this genus belongs not to the bacterial domain but the archaeal domain. Methanobacterium are nonmotile and live without oxygen, which is toxic to them, and they only inhabit anoxic environments.
Methanobrevibacter smithii is the predominant methanogenic archaeon in the microbiota of the human gut. M. smithii has a coccobacillus shape. It plays an important role in the efficient digestion of polysaccharides (complex sugars) by consuming the end products of bacterial fermentation (H2, CO2, acetate, and formate). M. smithii is a hydrogenotrophic methanogen that utilizes hydrogen by combining it with carbon dioxide to form methane. The removal of hydrogen by M. smithii is thought to allow an increase in the extraction of energy from nutrients by shifting bacterial fermentation to more oxidized end products.
Archaeol is a diether composed of two phytanyl chains linked to the sn-2 and sn-3 positions of glycerol. As its phosphate ester, it is a common component of the membranes of archaea.
Plagiopyla is a genus of ciliates. It includes nine species:
Karyorelictea is a class of ciliates in the subphylum Postciliodesmatophora. Most species are members of the microbenthos community, that is, microscopic organisms found in the marine interstitial habitat, though one genus, Loxodes, is found in freshwater.
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 undulating pattern than flagella. Cilia occur in all members of the group and are variously used in swimming, crawling, attachment, feeding, and sensation.
Colpidium colpoda are free-living ciliates commonly found in many freshwater environments including streams, rivers, lakes and ponds across the world. Colpidium colpoda is also frequently found inhabiting wastewater treatment plants. This species is used as an indicator of water quality and waste treatment plant performance.
Condylostoma is a genus of unicellular ciliate protists, belonging to the class Heterotrichea.
Kentrophoros is a genus of ciliates in the class Karyorelictea. Ciliates in this genus lack a distinct oral apparatus and depend primarily on symbiotic bacteria for their nutrition.
Licnophora is a genus of ciliates in the family Licnophoridae. They typically have an hourglass-like shape and live as ectocommensals on marine animals.
Metopus is a genus of anaerobic organisms from the family of Metopidae.
Parablepharismea is a class of free-living marine and brackish anaerobic ciliates that form a major clade of obligate anaerobes within the SAL group, together with the classes Muranotrichea and Armophorea.