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   » » Wiki: Trypanosomatida
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Trypanosomatida is a group of unicellular organisms distinguished by having only a single . The name is derived from the trypano (borer) and soma (body) because of the corkscrew-like motion of some trypanosomatid species. All members are exclusively , found primarily in . A few genera have life-cycles involving a secondary host, which may be a , or . These include several species that cause major diseases in humans. Some trypanosomatida are intracellular parasites, with the important exception of Trypanosoma brucei.


Medical importance
The three major diseases caused by trypanosomatids are; African (sleeping sickness, caused by Trypanosoma brucei and transmitted by ), South American trypanosomiasis (, caused by T. cruzi and transmitted by bugs), and (a set of trypanosomal diseases caused by various species of transmitted by
This review cites this research.
).


Evolution
The family is known from fossils of the extinct genus preserved in Burmese dating to the (100 mya) and from the (20–15 mya) of . The genus is also represented in Dominican amber in the extinct species T. antiquus.


Taxonomy
Three genera are (two hosts in the life cycle) – , and , while The remainder are monoxenous (one host in the life cycle). appears to be the first evolving branch in this order. Fifteen genera are recognised in the Trypanosomatidae and there are three subfamilies – , and . The genera in the subfamily are characterised by the presence of obligatory intracellular bacteria of the Kinetoplastibacterium genus.
:* Genus Gruby 1843
* Subfamily Votypka & Suková 2013
:* Genus Votypka & Suková 2013
* Subfamily sensu Maslov & Lukeš 2012
:* Clade Maslov & Lukeš 2012
::* Genus Léger 1902
::* Genus Kent 1880
::* Genus Schwarz 2015
:* Clade Maslov & Lukeš 2012
::* Genus Kostygov & Yurchenko 2017
::* Genus Mesnil & Brimont 1908
::* Genus Ross 1903
::* Genus Votýpka et al. 2015
::* Genus Cupolillo et al. 2000
::* Genus Shaw, Camargo et Teixeira 2016
* Subfamily Kostygov & Yurchenko 2015
:* Genus Kent 1880 non Donovan 1909
:* Genus Kostygov & Yurchenko 2015
:* Genus Donovan 1909
* Subfamily Votypka et al. 2014
:* Genus Souza & Corte-Real 1991
:* Genus Votypka et al. 2014
:* Genus Lwoff & Lwoff 1931


Life cycle
Some trypanosomatids only occupy a single host, while many others are : they live in more than one host species over their life cycle. This heteroxenous life cycle typically includes the of a bloodsucking and the blood and/or tissues of a . Rarer hosts include other bloodsucking invertebrates, such as , and other organisms such as . Different species go through a range of different morphologies at different stages of the life cycle, with most having at least two different morphologies. Typically the promastigote and epimastigote forms are found in insect hosts, trypomastigote forms in the mammalian and amastigotes in environments.

Among commonly studied examples, T. brucei, T. congolense, and T. vivax are extracellular, while T. cruzi and spp. are intracellular. Trypanosomatids with intracellular stages express proteins on their surfaces. de Paiva et al., 2015 illuminates δ-amastins' roles in intracellular success.


Sexual reproduction
Trypanosomatids that cause globally known diseases such ( species), African trypanosomiasis referred to as sleeping sickness ( Trypanosoma brucei), and ( Trypanosoma cruzi) were found to be capable of and genetic exchange. These findings indicate the capability for sexual reproduction in the Trypanosomatida.


Morphologies
A variety of different morphological forms appear in the life cycles of trypanosomatids, distinguished mainly by the position, length and the cell body attachment of the . The is found closely associated with the at the base of the flagellum and all species of trypanosomatid have a single nucleus. Most of these morphologies can be found as a life cycle stage in all trypanosomatid genera however certain morphologies are particularly common in a particular genus. The various morphologies were originally named from the genus where the morphology was commonly found, although this terminology is now rarely used because of potential confusion between morphologies and genus. Modern terminology is derived from the Greek; "mastig", meaning whip (referring to the flagellum), and a prefix which indicates the location of the flagellum on the cell. For example, the amastigote (prefix "a-", meaning no flagellum) form is also known as the leishmanial form as all have an amastigote life cycle stage.
  • (leishmanial). Amastigotes are a common morphology during an intracellular lifecycle stage in a mammalian host. All have an amastigote stage of the lifecycle. Leishmania amastigotes are particularly small and are among the smallest eukaryotic cells. The flagellum is very short, projecting only slightly beyond the flagellar pocket.
  • (leptomonad). The promastigote form is a common morphology in the insect host. The flagellum is found anterior of nucleus emerging directly from the anterior cell body. The kinetoplast is located in front of the nucleus, near the anterior end of the body.
  • (crithidial). Epimastigotes are a common form in the insect host and and , both parasites of insects, exhibit this form during their life cycles. The flagellum exits the cell anterior of nucleus and is connected to the cell body for part of its length by an undulating membrane. The kinetoplast is located between the nucleus and the anterior end.
  • (trypanosomal). This stage is characteristic of the genus in the mammalian host bloodstream as well as infective metacyclic stages in the fly vector. In trypomastigotes the is near the posterior end of the body, and the flagellum lies attached to the cell body for most of its length by an undulating membrane.
  • (herpetomonad). A rarer morphology where the flagellum posterior of nucleus, passing through a long groove in the cell.
  • . A morphotype where the flagellum does not extend beyond the deep flagellar pocket.

Image:LeishmaniaMexicana Amastigote SEM.jpg| Amastigote: False colour SEM micrograph of form Leishmania mexicana. The cell body is shown in orange and the flagellum is in red. 219 pixels/μm. Image:LeishmaniaMexicana Promastigote SEM.jpg| Promastigote: False colour SEM micrograph of form Leishmania mexicana. The cell body is shown in orange and the flagellum is in red. 119 pixels/μm. Image:TrypanosomaBrucei_ProcyclicTrypomastigote_SEM.jpg| Trypomastigote: False colour SEM micrograph of procyclic form Trypanosoma brucei. The cell body is shown in orange and the flagellum is in red. 84 pixels/μm.


Other features
Notable characteristics of trypanosomatids are the ability to perform of RNA and possession of , where much of their is confined to. The , another , was first identified in trypanosomes.


Bacterial endosymbiont
Six species of trypanosomatids are known to carry an additional proteobacterial , termed TPE (trypanosomatid proteobacterial endosymbionts). These trypansomatids ( Strigomonas oncopelti, S. culicis, S. galati, Angomonas desouzai, and ) are in turn known as SHTs, for symbiont-harboring trypanosomatids. All such symbionts have a shared evolutionary origin and are classified in the genus " Kinetoplastibacterium".

As with many symbionts, the bacteria have a much reduced genome compared to their free-living relatives of genera and . ( finds the genus sister to , a symbiont of .) Reflecting their inability to live alone, they have lost genes dedicated to essential biological functions, relying on the host instead. They have modified their division to become synchronized with the host. In S. culicis at least, the TPE helps the host by synthesizing and producing essential enzymes, staying tethered to the .

  • ( online). A comprehensive survey of the organisms' natural history.


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