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Trypanosomatida

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Trypanosomes
Temporal range: Albian to recent 100–0 Ma
Trypanosoma cruzi parasites
Scientific classification Edit this classification
Domain: Eukaryota
Phylum: Euglenozoa
Class: Kinetoplastea
Subclass: Metakinetoplastina
Order: Trypanosomatida
Kent 1880
Family: Trypanosomatidae
Doflein 1901
Subfamily

Trypanosomatida is a group of kinetoplastid excavates distinguished by having only a single flagellum. The name is derived from the Greek trypano (borer) and soma (body) because of the corkscrew-like motion of some trypanosomatid species. All members are exclusively parasitic, found primarily in insects.[1] A few genera have life-cycles involving a secondary host, which may be a vertebrate, invertebrate or plant. These include several species that cause major diseases in humans.[2] Trypanosomatida are intracellular parasites.

Medical importance

The three major human diseases caused by trypanosomatids are; African trypanosomiasis (sleeping sickness, caused by Trypanosoma brucei and transmitted by tsetse flies), South American trypanosomiasis (Chagas disease, caused by Trypanosoma cruzi and transmitted by triatomine bugs), and leishmaniasis (a set of trypanosomal diseases caused by various species of Leishmania transmitted by sandflies).

Evolution

The family is known from fossils of the extinct genus Paleoleishmania preserved in Burmese amber dating to the Albian (100 mya) and Dominican amber from the Burdigalian (20–15 mya) of Hispaniola.[3] The genus Trypanosoma is also represented in Dominican amber in the extinct species Trypanosoma antiquus.[4]

Taxonomy

Three genera are dixenous (two hosts in the life cycle) – Leishmania, Phytomonas and Trypanosoma and the remainder are monoxenous (one host in the life cycle). Paratrypanosoma appears to be the first evolving branch in this order. Fifteen genera are recognised in the Trypanosomatidae and there are three subfamilies – Blechomonadinae, Leishmaniinae and Strigomonadinae. The genera in the subfamily Strigomonadinae are characterised by the presence of obligatory intracellular bacteria of the Kinetoplastibacterium genus.

Life cycle

Some trypanosomatids only occupy a single host, while many others are heteroxenous: they live in more than one host species over their life cycle. This heteroxenous life cycle typically includes the intestine of a bloodsucking insect and the blood and/or tissues of a vertebrate. Rarer hosts include other bloodsucking invertebrates, such as leeches,[5] and other organisms such as plants. 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 bloodstream and amastigotes in intracellular environments.

Morphologies

Six main morphologies of trypanosomatids

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 flagellum. The kinetoplast is found closely associated with the basal body 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 Leishmania have an amastigote life cycle stage.

  • Amastigote (leishmanial).[6] Amastigotes are a common morphology during an intracellular lifecycle stage in a mammalian host. All Leishmania have an amastigote stage of the lifecycle. Leishmania amastigoes are particularly small and are among the smallest eukaryotic cells. The flagellum is very short, projecting only slightly beyond the flagellar pocket.
  • Promastigote (leptomonad).[6] The promastigote form is a common morphology in the insect host. The flagellum is found anterior of nucleus and flagellum not attached to the cell body. The kinetoplast is located in front of the nucleus, near the anterior end of the body.
  • Epimastigote (crithidial).[6] Epimastigotes are a common form in the insect host and Crithidia and Blastocrithidia, 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.
  • Trypomastigote (trypanosomal).[6] This stage is characteristic of the genus Trypanosoma in the mammalian host bloodstream as well as infective metacyclic stages in the fly vector. In trypomastigotes the kinetoplast 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.
  • Opisthomastigote (herpetomonad).[6] A rarer morphology where the flagellum posterior of nucleus, passing through a long groove in the cell.
  • Endomastigote.[7] A morphotype where the flagellum does not extend beyond the deep flagellar pocket.

Other features

Notable characteristics of trypanosomatids are the ability to perform trans-splicing of RNA and possession of glycosomes, where much of their glycolysis is confined to. The acidocalcisome, another organelle, was first identified in trypanosomes.[8]

References

  1. ^ Podlipaev S (May 2001). "The more insect trypanosomatids under study-the more diverse Trypanosomatidae appears". Int. J. Parasitol. 31 (5–6): 648–52. doi:10.1016/S0020-7519(01)00139-4. PMID 11334958.
  2. ^ Simpson AG, Stevens JR, Lukes J; Stevens; Lukes (April 2006). "The evolution and diversity of kinetoplastid flagellates". Trends Parasitol. 22 (4): 168–74. doi:10.1016/j.pt.2006.02.006. PMID 16504583.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  3. ^ Poinar, G. (2008). "Lutzomyia adiketis sp. n. (Diptera: Phlebotomidae), a vector of Paleoleishmania neotropicum sp. n. (Kinetoplastida: Trypanosomatidae) in Dominican amber". Parasites & Vectors. 1 (1): 22. doi:10.1186/1756-3305-1-22. PMC 2491605. PMID 18627624.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  4. ^ Poinar, G. (2005). "Triatoma dominicana sp. n. (Hemiptera: Reduviidae: Triatominae), and Trypanosoma antiquus sp. n. (Stercoraria: Trypanosomatidae), the First Fossil Evidence of a Triatomine-Trypanosomatid Vector Association". Vector-Borne and Zoonotic Diseases. 5 (1): 72–81. doi:10.1089/vbz.2005.5.72. PMID 15815152.
  5. ^ [1] "A new lineage of trypanosome from Australian vertebrates and terrestrial bloodsucking leeches (Haemadipsidae)"
  6. ^ a b c d e Hoare, Cecil A.; Wallace, Franklin G. (1966). "Developmental Stages of Trypanosomatid Flagellates: a New Terminology". Nature. 212 (5068): 1385–6. doi:10.1038/2121385a0. S2CID 4164112.
  7. ^ Merzlyak, Ekaterina; Yurchenko, Vyacheslav; Kolesnikov, Alexander A.; Alexandrov, Kirill; Podlipaev, Sergei A.; Maslov, Dmitri A. (2001-03-01). "Diversity and Phylogeny of Insect Trypanosomatids Based on Small Subunit rRNA Genes: Polyphyly of Leptomonas and Blastocrithidia". The Journal of Eukaryotic Microbiology. 48 (2): 161–169. doi:10.1111/j.1550-7408.2001.tb00298.x. ISSN 1066-5234. PMID 12095103. S2CID 13880469.
  8. ^ Docampo R, de Souza W, Miranda K, Rohloff P, Moreno SN; De Souza; Miranda; Rohloff; Moreno (March 2005). "Acidocalcisomes — conserved from bacteria to man". Nature Reviews Microbiology. 3 (3): 251–61. doi:10.1038/nrmicro1097. PMID 15738951. S2CID 31935658.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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