Kinetoplastid
{{short description|Flagellated protists belonging to the phylum Euglenozoa}}
{{automatic taxobox
| name = Kinetoplastida
| image = Trypanosoma_cruzi_crithidia.jpeg
| image_caption = Trypanosoma cruzi parasites
| taxon = Kinetoplastea
| authority =Honigberg, 1963 emend. Cavalier-Smith, 1981{{cite book|author=Honigberg, B. M.|year=1963|chapter=A contribution to systematics of the non-pigmented flagellates.|editor=Ludvík, J. |editor2=Lom, J. |editor3=Vávra J.|title=Progress in Protozoology: proceedings of the first International Congress on protozoology held at Prague|publisher=Academic Press}}Cavalier-Smith, T. (1981). Eukaryote kingdoms: seven or nine? Biosystems 14, 461–481.
| subdivision_ranks = Subdivisions
| subdivision =
- ?Bordnamonadidae
- ?Trypanophididae
- Prokinetoplastina
- Prokinetoplastida
- Metakinetoplastina
- Neobodonida
- Parabodonida
- Bodonida
- Trypanosomatida
| synonyms =
- Kinetoplastida Honigberg 1963 emend. Margulis 1974
}}
Kinetoplastida (or Kinetoplastea, as a class) is a group of flagellated protists belonging to the phylum Euglenozoa,{{cite book|last1=Berman|first1=Jules J.|title=Taxonomic Guide to Infectious Diseases: Understanding the Biologic Classes of Pathogenic Organisms|date=2012|publisher=Elsevier/Academic Press|location=London|isbn=978-0-12415-895-5|page=96|url=https://books.google.com/books?id=m7vT6i3xmC4C}}{{cite web|title=Kinetoplastida (kinetoplasts)|url=https://www.uniprot.org/taxonomy/5653|publisher=UniProt Consortium|access-date=22 January 2015}} and characterised by the presence of a distinctive organelle called the kinetoplast (hence the name), a granule containing a large mass of DNA. The group includes a number of parasites responsible for serious diseases in humans and other animals, as well as various forms found in soil and aquatic environments. The organisms are commonly referred to as "kinetoplastids" or "kinetoplasts".{{cite web|last=Lukes|first=Julius|title=Kinetoplastida|url=http://tolweb.org/Kinetoplastida/98013|publisher=The Tree of Life Web Project|access-date=10 September 2013|date=2009}}
The kinetoplastids were first defined by Bronislaw M. Honigberg in 1963 as the members of the flagellated protozoans. They are traditionally divided into the biflagellate Bodonidae and uniflagellate Trypanosomatidae; the former appears to be paraphyletic to the latter. One family of kinetoplastids, the trypanosomatids, is notable as it includes several genera which are exclusively parasitic. Bodo is a typical genus within kinetoplastida, which also includes various common free-living species which feed on bacteria. Others include Cryptobia and the parasitic Leishmania.
Taxonomy
=History=
Honigberg created the taxonomic names Kinetoplastida and Kinetoplastea in 1963. Since then there is no consensus on the use of either of the two as a definite taxon. Kinetoplastea is more widely used as the class,{{cite web|title=Kinetoplastea|url=http://eol.org/pages/2910504/overview|website=EOL|access-date=22 January 2015}}{{cite web|title=Kinetoplastea|url=http://www.marinespecies.org/aphia.php?p=taxdetails&id=562565|publisher=World Register of Marine Species|access-date=22 January 2015}}{{cite web|title=Kinetoplastea |url=http://zipcodezoo.com/key/protozoa/Kinetoplastea_Class.asp |publisher=ZipcodeZoo |access-date=22 January 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150122171006/http://zipcodezoo.com/key/protozoa/Kinetoplastea_Class.asp |archive-date=22 January 2015 }}{{cite web|title=Taxon: Class Kinetoplastea |url=http://taxonomicon.taxonomy.nl/TaxonName.aspx?id=198775&src=0 |publisher=The Taxonomicon |access-date=22 January 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150122151926/http://taxonomicon.taxonomy.nl/TaxonName.aspx?id=198775&src=0 |archive-date=22 January 2015 }}{{cite journal|last1=Moreira|first1=D.|title=An updated view of kinetoplastid phylogeny using environmental sequences and a closer outgroup: proposal for a new classification of the class Kinetoplastea|journal=International Journal of Systematic and Evolutionary Microbiology|date=2004|volume=54|issue=5|pages=1861–1875|doi=10.1099/ijs.0.63081-0|pmid=15388756|doi-access=free}} while Kinetoplastida is mostly used to designate the order,{{cite web|title=Kinetoplastida|url=http://eol.org/pages/5002/names?all=1|website=EOL|access-date=22 January 2015}}{{cite web|title=Kinetoplastida |url=http://www.metalife.com/NCBI%20Taxonomy/5653 |website=Metalife |access-date=22 January 2015 |url-status=dead |archive-url=https://web.archive.org/web/20150122153154/http://www.metalife.com/NCBI%20Taxonomy/5653 |archive-date=22 January 2015 }}{{cite web|title=Kinetoplastida|url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?lvl=0&id=5653|publisher=NCBI Taxonomy|access-date=22 January 2015}} but is also used as a class.{{cite book|last1=Gutierrez|first1=Yezid|title=Diagnostic Pathology of Parasitic Infections with Clinical Correlations|date=2000|publisher=Oxford University Press|location=New York|isbn=978-0-1951214-38|page=63|edition=2}} Lynn Margulis, who initially accepted Kinetoplastida as an order in 1974, later placed it as a class.{{cite book|first1=Lynn|last1=Margulis |first2=Michael J.|last2=Chapman |title=Kingdoms & Domains : An Illustrated Guide to the Phyla of Life on Earth|url=https://archive.org/details/fivekingdomsillu00marg_711|url-access=limited|year=2009|publisher=Academic Press/Elsevier|location=Amsterdam|isbn=978-0-12-373621-5|page=[https://archive.org/details/fivekingdomsillu00marg_711/page/n213 158]|edition=4}} Use of Kinetoplastida as an order also creates confusion as there is already an older name Trypanosomatida Kent, 1880, under which the kinetoplastids are most often placed.{{cite journal|last1=Deschamps|first1=P.|last2=Lara|first2=E.|last3=Marande|first3=W.|last4=Lopez-Garcia|first4=P.|last5=Ekelund|first5=F.|last6=Moreira|first6=D.|title=Phylogenomic analysis of kinetoplastids supports that trypanosomatids arose from within bodonids|journal=Molecular Biology and Evolution|year=2010|volume=28|issue=1|pages=53–58|doi=10.1093/molbev/msq289|pmid=21030427|doi-access=free}}
=Classification=
Kinetoplastida is divided into two subclasses - Metakinetoplastina and Prokinetoplastina.{{cite journal|last1=Moreira|first1=D|last2=López-García|first2=P|last3=Vickerman|first3=K|title=An updated view of kinetoplastid phylogeny using environmental sequences and a closer outgroup: proposal for a new classification of the class Kinetoplastea|journal=International Journal of Systematic and Evolutionary Microbiology|year=2004|volume=54|issue=Pt 5|pages=1861–1875|doi=10.1099/ijs.0.63081-0|pmid=15388756|doi-access=free}}{{cite journal|last1=Stevens|first1=JR|title=Kinetoplastid phylogenetics, with special reference to the evolution of parasitic trypanosomes|journal=Parasite|year= 2008|volume= 15|issue= 3|pages= 226–232|doi=10.1051/parasite/2008153226 |pmid=18814685|doi-access=free}}
- Family ?Bordnamonadidae Cavalier-Smith 2013
- Family ?Trypanophididae Poche 1911
- Subclass Prokinetoplastina Vickerman 2004
- Order Prokinetoplastida Vickerman 2004
- Family Ichthyobodonidae Isaksen et al. 2007
- Subclass Metakinetoplastina Vickerman 2004
- Order Neobodonida Vickerman 2004
- Family Rhynchomonadidae Cavalier-Smith 2016
- Family Neobodonidae Cavalier-Smith 2016
- Order Parabodonida Vickerman 2004
- Family Parabodonidae Cavalier-Smith 2016 [Cryptobiaceae Poche 1911; Cryptobiidae Vickerman 1976; Trypanoplasmatidae Hartmann & Chagas 1910]
- Order Bodonida Hollande 1952 emend. Vickerman 1976
- Family Bodonidae Bütschli 1883
- Order Trypanosomatida Kent 1880 stat. n. Hollande 1952 emend. Vickerman 2004
- Family Trypanosomatidae Doflein 1901 [Trypanomorphidae Woodcock 1906]
Morphology
Kinetoplastids are eukaryotic and possess normal eukaryotic organelles, for example the nucleus, mitochondrion, golgi apparatus and flagellum. Along with these universal structures, kinetoplastids have several distinguishing morphological features such as the kinetoplast, sub-pellicular microtubule array and paraflagellar rod.{{cn|date=March 2023}}
=Mitochondrion and kinetoplast DNA=
{{Main|Kinetoplast}}
The kinetoplast, after which the class is named, is a dense DNA-containing granule within the cell's single mitochondrion, containing many copies of the mitochondrial genome. The structure is made up of a network of concatenated circular DNA molecules and their related structural proteins along with DNA and RNA polymerases. The kinetoplast is found at the base of a cell's flagella and is associated to the flagellum basal body by a cytoskeletal structure.{{cn|date=March 2023}}
=Cytoskeleton=
The cytoskeleton of kinetoplastids is primarily made up of microtubules. These make a highly regular array, the sub-pellicular array, which runs parallel just under the cell surface along the long axis of the cell. Other microtubules with more specialised roles, such as the rootlet microtubules, are also present. Kinetoplastids are capable of forming actin microfilaments but their role in the cytoskeleton is not clear. Other cytoskeletal structures include the specialised attachment between the flagellum and the kinetoplast.{{cn|date=March 2023}}
=Flagella=
All kinetoplastids possess at least one flagellum; species in the order trypanosomatida have one and bodonida have two. In kinetoplastids with two flagella most forms have a leading and trailing flagellum, the latter of which may be attached to the side of the cell. The flagella are used for locomotion and attachment to surfaces. The bases of the flagella are found in a specialised pocket structure which is also the location of the cytostome.{{cn|date=March 2023}}
Flagellar membrane|
Flagellar axoneme|
Paraflagellar rod|
Flagellar attachment zone|
Flagellar-associated ER|
Pelicular microtubules|
Endosome, sorts material|
Glycosome|
Mitochondrion, creates ATP (energy) for the cell (discoid cristae)|
Golgi apparatus, modifies proteins and sends them out of the cell|
Flagellar pocket collar|
Flagellar pocket|
Connecting fibres|
Kinetoplast, DNA-containing granule|
Antipodal site}}]]
Life cycle
Kinetoplastids may be free-living or parasitic. The order trypanosomatida is notable as it includes many genera which are exclusively parasitic. Trypanosomatids may have simple life cycles in a single host or more complex ones which progress through multiple differentiation stages in two hosts. Dramatic morphological changes are possible between lifecycle stages. Diseases caused by members of the order trypanosomatida include sleeping sickness and Chagas disease, caused by species of Trypanosoma, and leishmaniasis, caused by species of Leishmania.{{cite journal|vauthors=Scheckenbach F, Wylezich C, Mylnikov AP, Weitere M, Arndt H|title=Molecular comparisons of freshwater and marine isolates of the same morphospecies of heterotrophic flagellates|journal=Appl Environ Microbiol|date=October 2006|volume=72|issue=10|pages=6638–6643|doi=10.1128/AEM.02547-05|pmid=17021215|pmc=1610283|bibcode=2006ApEnM..72.6638S }}
Trypanosoma brucei can undergo meiosis as a likely part of a sexual cycle.{{cite journal |vauthors=Peacock L, Ferris V, Sharma R, Sunter J, Bailey M, Carrington M, Gibson W |title=Identification of the meiotic life cycle stage of Trypanosoma brucei in the tsetse fly |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=108 |issue=9 |pages=3671–6 |year=2011 |pmid=21321215 |pmc=3048101 |doi=10.1073/pnas.1019423108 |url=http://psasir.upm.edu.my/25359/1/Identification%20of%20the%20meiotic%20life%20cycle%20stage%20of%20Trypanosoma%20brucei%20in%20the%20tsetse%20fly.pdf|bibcode=2011PNAS..108.3671P |doi-access=free }}{{cite journal |vauthors=Gibson W |title=Liaisons dangereuses: sexual recombination among pathogenic trypanosomes |journal=Res. Microbiol. |volume=166 |issue=6 |pages=459–66 |year=2015 |pmid=26027775 |doi=10.1016/j.resmic.2015.05.005 |url=https://research-information.bristol.ac.uk/en/publications/liaisons-dangereuses(1ecb5cba-da25-4e93-a3cb-b00a0477cb23).html|hdl=1983/1ecb5cba-da25-4e93-a3cb-b00a0477cb23 |hdl-access=free }} Leishmania major is also capable of a meiotic process that is likely part of a sexual cycle.{{cite journal |vauthors=Akopyants NS, Kimblin N, Secundino N, Patrick R, Peters N, Lawyer P, Dobson DE, Beverley SM, Sacks DL |title=Demonstration of genetic exchange during cyclical development of Leishmania in the sand fly vector |journal=Science |volume=324 |issue=5924 |pages=265–8 |year=2009 |pmid=19359589 |pmc=2729066 |doi=10.1126/science.1169464 |bibcode=2009Sci...324..265A }}
Gallery
Cryptobia helicis x15,000 Scanning EM.jpg|Cryptobia sp.
File:Bodo saltans - 400x (13895749563).jpg|Bodo sp.
File:Trypanosoma sp. PHIL 613 lores.jpg|Trypanosoma sp.
References
Bibliography
- Lumsden, W.H.R. & D.A. Evans (eds.). 1976-1979. Biology of the Kinetoplastida, 2 vols. London: Academic Press.