The OPISTHOKONTS (Greek : ὀπίσθιος (_opísthios_) = "rear, posterior" + κοντός (_kontós_) = "pole" i.e. "flagellum") are a broad group of eukaryotes , including both the animal and fungus kingdoms , together with the eukaryotic microorganisms that are sometimes grouped in the paraphyletic phylum Choanozoa (conventionally assigned to the protist "kingdom"). The opisthokonts, sometimes referred to as the "Fungi/ Metazoa group", are generally recognized as a monophyletic clade .
* 1 Flagella * 2 History * 3 Taxonomy * 4 Gallery * 5 References * 6 External links
One common characteristic of opisthokonts is that flagellate cells, such as the sperm of most animals and the spores of the chytrid fungi , propel themselves with a single _posterior_ flagellum. It is this feature that gives the group its name. In contrast, flagellate cells in other eukaryote groups propel themselves with one or more _anterior_ flagella (see Zoospore morphological types ). In some opisthokont groups, including most of the fungi, flagellate cells have been lost.
The close relationship between animals and fungi was suggested by Thomas Cavalier-Smith in 1987, who used the informal name opisthokonta (the formal name has been used for the chytrids by Copeland in 1956), and was supported by later genetic studies.
Early phylogenies placed fungi near the plants and other groups that have mitochondria with flat cristae , but this character varies. More recently, it has been said that holozoa (animals) and holomycota (fungi) are much more closely related to each other than either is to plants, because opisthokonts have a triple fusion of carbamoyl phosphate synthetase , dihydroorotase , and aspartate carbamoyltransferase that is not present in plants, and plants have a fusion of thymidylate synthase and dihydrofolate reductase not present in the opisthokonts. Animals and fungi are also more closely related to amoebas than they are to plants, and plants are more closely related to the SAR supergroup of protists than they are to animals or fungi. Animals and fungi are both heterotrophs , unlike plants, and while fungi are sessile like plants, there are also sessile animals.
Cavalier-Smith and Stechmann argue that the uniciliate eukaryotes such as opisthokonts and Amoebozoa , collectively called unikonts , split off from the other biciliate eukaryotes, called bikonts , shortly after they evolved .
One hypothesis of eukaryote relationships
Opisthokonts are divided into Holomycota or Nucletmycea (fungi and all organisms more closely related to fungi than to animals) and Holozoa (animals and all organisms more closely related to animals than to fungi); no opisthokonts basal to the Holomycota/ Holozoa split have yet been identified. However, a 2013 study suggests that there may be many more unicellular opisthokonts than previously suspected. Holomycota and Holozoa are composed of the following groups.
* including chytrids (previously thought to be protists ) * including microsporidia (previously thought to be protists) * including _Hyaloraphidium _ (previously thought to be a green alga , now thought to be a chytrid) * excluding oomycetes (water molds) (now thought to be stramenopiles ) * excluding labyrinthulomycetes (slime nets) (now thought to be stramenopiles ) * excluding myxomycetes (now thought to be amoebozoans )
* Capsasporida (capsasporid amoebae) * Ministeriida (ministeriid amoebae)
* including Myxozoa (previously thought to be protists and now understood to be cnidarians )
The paraphyletic taxon Choanozoa includes either non-animal holozoans, or non-animal, non-fungi opisthokonts.
The choanoflagellates have a circular mitochondrial DNA genome with long intergenic regions. This is four times as large as animal mitochondrial genomes and contains twice as many protein genes.
_ Corallochytrium _ seem likely to be more closely related to the fungi than to the animals on the basis of the presence of ergosterol in their membranes and being capable of synthesis of lysine via the α-aminoadipate (AAA) pathway .
The ichthyosporeans have a two amino acid deletion in their elongation factor 1 α gene that is considered characteristic of fungi.
The ichthyosl genome is >200 kilobase pairs in length and consists of several hundred linear chromosomes that share elaborate terminal-specific sequence patterns.
One view of the great kingdoms and their stem groups.
Microsporidian spore ( Microsporidia ) *
_Sphaeroforma _ sp. ( Mesomycetozoea ) *
_Desmarella _ sp. colony ( Choanoflagellatea ) *
* ^ Parfrey, Laura Wegener; Lahr, Daniel J. G.; Knoll, Andrew H.; Katz, Laura A. (August 16, 2011). "Estimating the timing of early eukaryotic diversification with multigene molecular clocks". _Proceedings of the National Academy of Sciences of the United States of America_. 108 (33): 13624–13629. PMC 3158185 _. PMID 21810989 . doi :10.1073/pnas.1110633108 . * ^ Copeland, H. F. (1956). The Classification of Lower Organisms_. Palo Alto: Pacific Books. * ^ _A_ _B_ Cavalier-Smith, T. (1987). "The origin of fungi and pseudofungi". In Rayner, Alan D. M. (ed.). _Evolutionary biology of Fungi_. Cambridge: Cambridge Univ. Press. pp. 339–353. ISBN 0-521-33050-5 . CS1 maint: Extra text: editors list (link ) * ^ Adl, S.M.; et al. (September–October 2005). "The new higher level classification of eukaryotes with emphasis on the taxonomy of protists". _Journal of eukaryotic microbiology_. 52: 399–451. PMID 16248873 . doi :10.1111/j.1550-7408.2005.00053.x . * ^ Shalchian-Tabrizi K, Minge MA, Espelund M, et al. (7 May 2008). Aramayo R, ed. "Multigene phylogeny of choanozoa and the origin of animals". _PLoS ONE_. 3 (5): e2098. Bibcode :2008PLoSO...3.2098S. PMC 2346548 _. PMID 18461162 . doi :10.1371/journal.pone.0002098 . * ^ A_ _B_ _C_ Steenkamp ET, Wright J, Baldauf SL (January 2006). "The protistan origins of animals and fungi". _Mol. Biol. Evol_. 23 (1): 93–106. PMID 16151185 . doi :10.1093/molbev/msj011 . * ^ "Fungi/ Metazoa group". Retrieved 2009-03-08. * ^ Huang, Jinling; Xu, Ying; Gogarten, Johann Peter (November 2005). "The presence of a haloarchaeal type tyrosyl-tRNA synthetase marks the opisthokonts as monophyletic". _Molecular Biology and Evolution_. 22 (11): 2142–2146. PMID 16049196 . doi :10.1093/molbev/msi221 . * ^ Parfrey, Laura Wegener; et al. (December 2006). "Evaluating support for the current classification of eukaryotic diversity" . _PLOS Genetics_. 2 (12): e220. PMC 1713255 _. PMID 17194223 . doi :10.1371/journal.pgen.0020220 . * ^ Torruella, Guifré; et al. (February 2012). "Phylogenetic relationships within the Opisthokonta based on phylogenomic analyses of conserved single-copy protein domains" . Molecular Biology and Evolution_. 29 (2): 531–544. PMC 3350318 _. PMID 21771718 . doi :10.1093/molbev/msr185 . * ^ Wainright PO, Hinkle G, Sogin ML, Stickel SK (April 1993). " Monophyletic origins of the metazoa: an evolutionary link with fungi". Science_. 260 (5106): 340–2. Bibcode :1993Sci...260..340W. PMID 8469985 . doi :10.1126/science.8469985 . * ^ Stechmann, A.; Cavalier-Smith, T. (5 July 2002). "Rooting the eukaryote tree by using a derived gene fusion". _Science_. 297 (5578): 89–91. Bibcode :2002Sci...297...89S. PMID 12098695 . doi :10.1126/science.1071196 . * ^ Del Campo, J.; Ruiz-Trillo, I. (April 2013). "Environmental survey meta-analysis reveals hidden diversity among unicellular opisthokonts" . _Molecular Biology and Evolution_. 30 (4): 802–805. PMC 3603316 _. PMID 23329685 . doi :10.1093/molbev/mst006 .
* ^ Phylogeny based on:
* Eichinger, L.; Pachebat, J. A.; Glöckner, G.; Rajandream, M. A.; Sucgang, R.; Berriman, M.; Song, J.; Olsen, R.; et al. (2005). "The genome of the social amoeba Dictyostelium discoideum_" . _Nature_. 435 (7038): 43–57. PMC 1352341 _. PMID 15875012 . doi :10.1038/nature03481 . * Steenkamp, E. T.; Wright, J.; Baldauf, S. L. (2005). "The Protistan Origins of Animals and Fungi". Molecular Biology and Evolution_. 23 (1): 93–106. PMID 16151185 . doi :10.1093/molbev/msj011 .
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