Trypanosomatida is a group of kinetoplastid unicellular organisms 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. 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. Some trypanosomatida are intracellular parasites, with the important exception of Trypanosoma brucei.
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 T. 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. The genus Trypanosoma is also represented in Dominican amber in the extinct species T. antiquus. The family contains both dixenous (two hosts in the life cycle) and monoxenous (one host in the life cycle) species. The dixenous lifestyle occurs in three lineages: one for plants (Phytomonas), two for vertebrates (Leishmania/Porcisia/Endotrypanum and Trypanosoma). These are believed to have evolved separately. The rest of the family is monoxenous, with the most common host being insects. The trypanosomatids are also notable for the presence of a glycosome, which contains sugar-breakdown enzymes that are most closely related to alage. Some researchers believe that the presence of an entire block of enzymes is indicative of a past where an ancestral trypanosomatid engulfed an alga (probably a green alga) and co-opted some of its genetic material. Regardless of the cause of this occurrence, this difference in sugar metabolism is a potential target of therapies.
Taxonomy Family Trypanosomatidae Calkins 1926 [Trypanomorphidae Woodcock 1906; Trypanosomataceae Senn 1911] Genus Agamomonas Grassé 1952 Genus Batracoleishmania Dasgupta 2011 Genus Blastocrithidia Laird 1959 Genus Cercoplasma Roubaud 1911 Genus Cystotrypanosoma Roubaud 1911 Genus Jaenimonas Votypka & Hamilton 2015 Genus Lamellasoma Davis 1947 Genus Leptowallaceina Podlipaev & Frolov 2000 Genus Lewisonella Chalmers 1918 nomen dubium Genus Malacozoomonas Nicoli, Penaud & Timon-David 1972 Genus Nematodomonas Nicoli, Penaud & Timon-David 1972 Genus †Paleoleishmania Poinar & Poinar, 2004 Genus †Paleotrypanosoma Poinar 2008 Genus Paramecioides Grassé 1882 Genus Sauroleishmania Ranque 1973 Genus Sergeia Svobodová et al. 2007 non Stimpson 1860 non Nasonov 1923 non Sergio Manning & Lemaitre 1994 Genus Trypanomonas Danilewsky 1885 Genus Trypanomorpha Woodcock 1906 Genus Undulina Lankester 187 Genus Wallaceina Bulat, Mokrousov & Podlipaev 1999 [Proteomonas Podlipaev, Frolov & Kolesnikov 1990 non Hill & Wetherbee 1986] Genus Wallacemonas Kostygov & Yurchenko 2014 Subfamily Paratrypanosomatinae Votýpka & Lukeš 2013 Genus Paratrypanosoma Votypka & Lukes 2013 Subfamily Trypanosomatinae Genus Trypanosoma Gruby 1843 Subfamily Blechomonadinae Votypka & Suková 2013 Genus Blechomonas Votypka & Suková 2013 Subfamily Leishmaniinae sensu Maslov & Lukeš 2012 Infrafamily Crithidiatae Kostygov & Yurchenko 2017 Genus Crithidia Léger 1902 Genus Leptomonas Kent 1880 Genus Lotmaria Schwarz 2015 Infrafamily Leishmaniatae Maslov & Lukeš 2012 Genus Borovskyia Kostygov & Yurchenko 2017 Genus Endotrypanum Mesnil & Brimont 1908 Genus Leishmania Ross 1903 Genus Novymonas Votýpka et al. 2015 Genus Paraleishmania Cupolillo et al. 2000 Genus Zelonia Shaw, Camargo et Teixeira 2016 Subfamily Phytomonadinae Kostygov & Yurchenko 2015 Genus Herpetomonas Kent 1880 non Donovan 1909 Genus Lafontella Kostygov & Yurchenko 2015 Genus Phytomonas Donovan 1909 Subfamily Strigomonadinae Votypka et al. 2014 – characterised by the presence of obligatory intracellular bacteria of the Kinetoplastibacterium genus. Genus Angomonas Souza & Corte-Real 1991 Genus Kentomonas Votypka et al. 2014 Genus Strigomonas Lwoff & Lwoff 1931
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, 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. Among commonly studied examples, T. brucei, T. congolense, and T. vivax are extracellular, while T. cruzi and Leishmania spp. are intracellular. Trypanosomatids with intracellular stages express δ-amastin 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 leishmaniasis (Leishmania species), African trypanosomiasis referred to as sleeping sickness (Trypanosoma brucei), and Chagas disease (Trypanosoma cruzi) were found to be capable of meiosis and genetic exchange. These findings indicate the capability for sexual reproduction in the Trypanosomatida.
Morphologies
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