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Labyrinthulomycetes
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<h2 id="characteristics">Characteristics</h2> <p>Although they are outside the cells, the filaments of Labyrinthulomycetes are surrounded by a <a href="/facts/Cell_membrane/vAXSD0DT">membrane</a>. They are formed and connected with the cytoplasm by a unique organelle called a sagenogen or bothrosome. The cells are uninucleated and typically ovoid, and move back and forth along the <a href="/facts/Amorphous/YTIzmrR0">amorphous</a> network at speeds varying from 5-150 μm per minute. Among the labyrinthulids, the cells are enclosed within the tubes, and among the thraustochytrids, they are attached to their sides. </p> <h2 id="evolution">Evolution</h2> <h3>Evolutionary origin</h3> <p>Labyrinthulomycetes are not <a href="/facts/Fungi/BHo5DP59">fungi</a>, but a <a href="/facts/Monophyletic/pD8Yrrn6">monophyletic</a> group of <a href="/facts/Eukaryote/SkwyPLMA">eukaryotes</a> within the <a href="/facts/Stramenopiles/6zmwHPEU">Stramenopiles</a>. They belong to the phylum <a href="/facts/Bigyra/DQKsDcry">Bigyra</a>, which contains other heterotrophic microorganisms such as the <a href="/facts/Bicosoecid/JaeIDHYV">bicosoecids</a>. Considering that the <a href="/facts/Stramenopiles/6zmwHPEU">plastids</a> from <a href="/facts/Stramenopiles/6zmwHPEU">Stramenopiles</a> are possibly the result of an event of <a href="/facts/Endosymbiosis/eWvSS8ID">endosymbiosis</a> in their <a href="/facts/Last_common_ancestor/Cbg1YLCV">last common ancestor</a>, the bicosoecids and the labyrinthulomycetes could have originated from a <a href="/facts/Mixotrophic/S7ePtbLm">mixotrophic</a> <a href="/facts/Algal/U1ftVKU3">algal</a> common ancestor that <a href="/facts/Secondary_loss/t21GWRl1">secondarily lost</a> their plastids.<a class="footnote-ref" id="fnref:6" href="#fn:6"><sup>6</sup></a> </p><p>Some characteristics of the labyrinthulomycetes can be explained by their origin from ancestral plastids. They produce <a href="/facts/Omega-3_fatty_acid/X72Ynqov">omega-3 poly-unsaturated fatty acids</a> using a <a href="/facts/Desaturase/edjFHcuj">desaturase</a> usually present in <a href="/facts/Chloroplast/fo8U3yTx">chloroplasts</a>. The zoospores of <a href="/facts/Labyrinthulid/bD1RHSwc">labyrinthulids</a> have an <a href="/facts/Eyespot_apparatus/61RsMDLr">eyespot</a> composed of membrane-bound granules that resembles eyespots of photosynthetic stramenopiles, which are either within a plastid or believed to be derived from a plastid.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> </p><p>Within <a href="/facts/Bigyra/DQKsDcry">Bigyra</a>, the labyrinthulomycetes are the <a href="/facts/Sister_group/pNNuVex4">sister group</a> to <a href="/facts/Eogyrea/l5PpEUVM">Eogyrea</a>, a class containing the species <i>Pseudophyllomitus vesiculosus</i> and the <a href="/facts/Environmental_DNA/GG7dGwyP">environmental</a> clade called MAST-4. Together they compose the subphylum <a href="/facts/Sagenista/l5PpEUVM">Sagenista</a>.<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a><a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a> </p> <table><tbody><tr><td><a href="/facts/Stramenopiles/6zmwHPEU">Stramenopiles</a></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td><a href="/facts/Gyrista/lSn1g6V1">Gyrista</a></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><a href="/facts/Ochrophyta/1kduBajE">Ochrophyta</a> </p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Pseudofungi/ltXLY2qV">Pseudofungi</a> </p></td></tr><tr><td><i>plastid loss</i></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td><a href="/facts/Bigyra/DQKsDcry">Bigyra</a></td><td rowspan="2"><table><tbody><tr><td><a href="/facts/Opalozoa/Vro8HH61">Opalozoa</a></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><a href="/facts/Placidozoa/YrhJwmQL">Placidozoa</a> </p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Bicosoecida/JaeIDHYV">Bicosoecida</a> </p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td><a href="/facts/Sagenista/l5PpEUVM">Sagenista</a></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>Labyrinthulomycetes </p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Eogyrea/l5PpEUVM">Eogyrea</a></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td><i>plastid loss</i></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Platysulcea/WsxAraMH">Platysulcea</a></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table> <h2 id="classification">Classification</h2> <p>Labyrinthulomycetes or Labyrinthulea used to compose the defunct fungal <a href="/facts/Phylum/ELT3Jyax">phylum</a> Labyrinthulomycota.<a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a> They were originally considered unusual <a href="/facts/Slime_mould/bA6b9Vss">slime moulds</a>, although they are not very similar to the other sorts. The structure of their <a href="/facts/Zoospores/TjpX7rnQ">zoospores</a> and genetic studies show them to be a primitive group of <a href="/facts/Heterokont/WFevSvg2">heterokonts</a>, but their classification and treatment remains somewhat unsettled. </p><p>This class usually contained two orders, Labyrinthulales and Thraustochytriales (ICBN), or <a href="/facts/Labyrinthulida/bD1RHSwc">Labyrinthulida</a> and <a href="/facts/Thraustochytrida/G17rGlfo">Thraustochytrida</a> (ICZN), but a different classification has recently been proposed.<a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a><a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a><a class="footnote-ref" id="fnref:13" href="#fn:13"><sup>13</sup></a><a class="footnote-ref" id="fnref:14" href="#fn:14"><sup>14</sup></a><a class="footnote-ref" id="fnref:15" href="#fn:15"><sup>15</sup></a> </p> <ul><li>Order Labyrinthulales/<a href="/facts/Labyrinthulida/bD1RHSwc">Labyrinthulida</a> E. A. Bessey 1950/Doffein 1901 <ul><li>Family Aplanochytriaceae/Aplanochytriidae Leander ex Cavalier-Smith 2012 <ul><li><i><a href="/facts/Aplanochytrium/xbGyRWdd">Aplanochytrium</a></i> Bahnweg & Sparrow 1972 [=<i>Labyrinthuloides</i> Perkins 1973]</li></ul></li> <li>Family Labyrinthulaceae/Labyrinthulidae Haeckel 1868/Cinekowksa 1867 <ul><li><i>Labyrinthomyxa</i> Duboscq 1921</li> <li><i><a href="/facts/Pseudoplasmodium/K3yRSUjh">Pseudoplasmodium</a></i> Molisch 1925</li> <li><i><a href="/facts/Labyrinthula/n6F3o8uX">Labyrinthula</a></i> Cienkowski 1864 [=<i>Labyrinthodictyon</i> Valkanov 1969; <i>Labyrinthorhiza</i> Chadefaud 1956]</li></ul></li> <li>Family-level clade "Stellarchytriaceae/Stellarchytriidae" – this group is provisionally placed in <a href="/facts/Labyrinthulida/bD1RHSwc">Labyrinthulida</a><a class="footnote-ref" id="fnref:16" href="#fn:16"><sup>16</sup></a><a class="footnote-ref" id="fnref:17" href="#fn:17"><sup>17</sup></a> but, according to phylogenetic analyses, diverges before the rest of labyrinthulean clades.<a class="footnote-ref" id="fnref:18" href="#fn:18"><sup>18</sup></a> <ul><li><i>Stellarchytrium</i> FioRito & Leander 2016</li></ul></li></ul></li> <li>Order Oblongichytriales/Oblongichytrida <ul><li>Family Oblongichytriaceae/Oblongichytriidae Cavalier-Smith 2012 <ul><li><i>Oblongichytrium</i> Yokoyama & Honda 2007</li></ul></li></ul></li> <li>Order <a href="/facts/Thraustochytriales/G17rGlfo">Thraustochytriales</a>/<a href="/facts/Thraustochytrida/G17rGlfo">Thraustochytrida</a> Sparrow 1973 <ul><li><i>Pyrrhosorus</i> Juel 1901</li> <li><i>Thanatostrea</i> Franc & Arvy 1969</li> <li>Family Althornidiaceae/Althorniidae Jones & Alderman 1972 <ul><li><i>Althornia</i> Jones & Alderman 1972</li></ul></li> <li>Family Thraustochytriacae/<a href="/facts/Thraustochytriidae/Y27fQ3Un">Thraustochytriidae</a> Sparrow ex Cejp 1959 <ul><li><i>Japanochytrium</i> Kobayasi & Ôkubo 1953</li> <li><i>Monorhizochytrium</i> Doi & Honda 2017</li> <li><i>Sicyoidochytrium</i> Yokoy., Salleh & Honda 2007</li> <li><i>Aurantiochytrium</i> Yokoy. & Honda 2007</li> <li><i>Ulkenia</i> Gaertn. 1977</li> <li><i>Parietichytrium</i> Yokoy., Salleh & Honda 2007</li> <li><i>Botryochytrium</i> Yokoy., Salleh & Honda 2007</li> <li><i><a href="/facts/Schizochytrium/LTEK4xjs">Schizochytrium</a></i> Goldst. & Belsky emend. Booth & Mill.</li> <li><i>Thraustochytrium</i> Sparrow 1936</li> <li><i>Hondaea</i> Amato & Cagnac 2018</li> <li><i>Labyrinthulochytrium</i> Hassett & Gradinger 2018<i><a class="footnote-ref" id="fnref:19" href="#fn:19"><sup>19</sup></a></i></li></ul></li></ul></li> <li>Order "Amphitremidales"/<a href="/facts/Amphitremida/bD1RHSwc">Amphitremida</a> Gomaa et al. 2013 <ul><li>Family "Amphitremidiaceae"/Amphitremidae Poch 1913 <ul><li><i>Paramphitrema</i> Valkanov 1970</li> <li><i>Archerella</i> Loeblich & Tappan 1961</li> <li><i><a href="/facts/Amphitrema/85Uz6EYl">Amphitrema</a></i> Archer 1867</li></ul></li> <li>Family "Diplophrydaceae"/Diplophryidae Cavalier-Smith 2012 <ul><li><i>Diplophrys</i> Barker 1868</li></ul></li></ul></li> <li>Order "Amphifilales"/Amphifilida Cavalier Smith 2012 <ul><li>Family Sorodiplophryidae Cavalier-Smith 2012 <ul><li><i>Sorodiplophrys</i> Olive & Dykstra 1975</li> <li><i>Fibrophrys</i> Takahashi et al. 2016</li></ul></li> <li>Family Amphifilidae Cavalier-Smith 2012 <ul><li>Genus <i>Amphifila</i> Cavalier-Smith 2012</li></ul></li></ul></li></ul> <h2 id="genetic-code">Genetic code</h2> <p>The labyrinthulomycete <i>Thraustochytrium aureum</i> is notable for the <a href="/facts/Genetic_code/zUK0bY1B">alternative genetic code</a> of its <a href="/facts/Mitochondria/ErHqrdJ1">mitochondria</a> which use TTA as a <a href="/facts/Stop_codon/1yuAkq9X">stop codon</a> instead of coding for <a href="/facts/Leucine/X7YrkvA5">Leucine</a>.<a class="footnote-ref" id="fnref:20" href="#fn:20"><sup>20</sup></a> This code is represented by <a href="/facts/National_Center_for_Biotechnology_Information/6Xj5wCbu">NCBI</a> translation table 23, <a href="/facts/Thraustochytrium_mitochondrial_code/Q2DT7Ytm"><i>Thraustochytrium</i> mitochondrial code</a>.<a class="footnote-ref" id="fnref:21" href="#fn:21"><sup>21</sup></a> </p> <table><tbody><tr><th>Genetic code</th><th>Translation table</th><th>DNA codon</th><th>RNA codon</th><th colspan="3">Translation with this code</th><th></th><th><a href="/facts/DNA_codon_table/9ceoWs0X">Standard code</a>(Translation table 1)</th></tr><tr><td rowspan="1"><a href="/facts/Thraustochytrium_mitochondrial_code/Q2DT7Ytm"><i>Thraustochytrium</i> mitochondrial</a></td><td rowspan="1">23</td><td>TTA</td><td>UUA</td><td colspan="3">STOP = Ter (*)</td><td></td><td>Leu (L)</td></tr></tbody></table> <h2 id="gallery">Gallery</h2> <h2 id="external-links">External links</h2> <a href="/facts/Wikisource/TrMdylNf">Wikisource</a> has the text of the <a href="/facts/Encyclop%25C3%25A6dia_Britannica_Eleventh_Edition/mPElbCRY">1911 <i>Encyclopædia Britannica</i></a> article "Labyrinthulidea". <ul><li><a href="https://web.archive.org/web/20070417091256/http://www.botany.uga.edu/zoosporicfungi/labyrint.htm">Labyrinthulomycota</a></li></ul> <a href="/facts/Wikispecies/b2GhA7Le">Wikispecies</a> has information related to <i>Labyrinthulomycetes</i>. <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Cavalier-Smith, T. (1997). "Sagenista and bigyra, two phyla of heterotrophic heterokont chromists". Archiv für Protistenkunde. 148 (3): 253–267. doi:10.1016/S0003-9365(97)80006-1. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Tsui, Clement K M; Marshall, Wyth; Yokoyama, Rinka; Honda, Daiske; Lippmeier, J Casey; Craven, Kelly D; Peterson, Paul D; Berbee, Mary L (January 2009). "Labyrinthulomycetes phylogeny and its implications for the evolutionary loss of chloroplasts and gain of ectoplasmic gliding". Molecular Phylogenetics and Evolution. 50 (1): 129–40. doi:10.1016/j.ympev.2008.09.027. PMID 18977305. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>Schärer, L.; Knoflach, D.; Vizoso, D. B.; Rieger, G.; Peintner, U. (2007). "Thraustochytrids as novel parasitic protists of marine free-living flatworms: Thraustochytrium caudivorum sp. nov. Parasitizes Macrostomum lignano" (PDF). Marine Biology. 152 (5): 1095. doi:10.1007/s00227-007-0755-4. S2CID 4836350. <a href="http://doc.rero.ch/record/319983/files/227_2007_Article_755.pdf" target="_blank">http://doc.rero.ch/record/319983/files/227_2007_Article_755.pdf</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>Pan, Jingwen (2016). Labyrinthulomycetes diversity meta-analysis (MSc). University of British Columbia. doi:10.14288/1.0223199. <a href="https://open.library.ubc.ca/media/stream/pdf/24/1.0223199/3" target="_blank">https://open.library.ubc.ca/media/stream/pdf/24/1.0223199/3</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>Gomaa, Fatma; Mitchell, Edward A. D.; Lara, Enrique (2013). "Amphitremida (poche, 1913) is a new major, ubiquitous labyrinthulomycete clade". PLoS One. 8 (1): e53046. Bibcode:2013PLoSO...853046G. doi:10.1371/journal.pone.0053046. PMC 3544814. PMID 23341921. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3544814" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3544814</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></p></li> <li id="fn:6"><p>Tsui, Clement K M; Marshall, Wyth; Yokoyama, Rinka; Honda, Daiske; Lippmeier, J Casey; Craven, Kelly D; Peterson, Paul D; Berbee, Mary L (January 2009). "Labyrinthulomycetes phylogeny and its implications for the evolutionary loss of chloroplasts and gain of ectoplasmic gliding". Molecular Phylogenetics and Evolution. 50 (1): 129–40. doi:10.1016/j.ympev.2008.09.027. PMID 18977305. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></p></li> <li id="fn:7"><p>Tsui, Clement K M; Marshall, Wyth; Yokoyama, Rinka; Honda, Daiske; Lippmeier, J Casey; Craven, Kelly D; Peterson, Paul D; Berbee, Mary L (January 2009). "Labyrinthulomycetes phylogeny and its implications for the evolutionary loss of chloroplasts and gain of ectoplasmic gliding". Molecular Phylogenetics and Evolution. 50 (1): 129–40. doi:10.1016/j.ympev.2008.09.027. PMID 18977305. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:7" class="footnote-back-ref">↩</a></p></li> <li id="fn:8"><p>Cavalier-Smith, Thomas (2017). "Kingdom Chromista and its eight phyla: a new synthesis emphasising periplastid protein targeting, cytoskeletal and periplastid evolution, and ancient divergences". Protoplasma. 255 (1): 297–357. doi:10.1007/s00709-017-1147-3. PMC 5756292. PMID 28875267. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5756292" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5756292</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></p></li> <li id="fn:9"><p>Thakur, Rabindra; Shiratori, Takashi; Ishida, Ken-ichiro (2019). "Taxon-rich Multigene Phylogenetic Analyses Resolve the Phylogenetic Relationship Among Deep-branching Stramenopiles". Protist. 170 (5): 125682. doi:10.1016/j.protis.2019.125682. ISSN 1434-4610. PMID 31568885. S2CID 202865459. <a href="https://www.sciencedirect.com/science/article/pii/S1434461018300865" target="_blank">https://www.sciencedirect.com/science/article/pii/S1434461018300865</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>Bennett, Reuel M.; Honda, D.; Beakes, Gordon W.; Thines, Marco (2017). "Chapter 14. Labyrinthulomycota". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Springer. pp. 507–542. doi:10.1007/978-3-319-28149-0_25. ISBN 978-3-319-28147-6. <a href="978-3-319-28147-6" target="_blank">978-3-319-28147-6</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>Gomaa, Fatma; Mitchell, Edward A. D.; Lara, Enrique (2013). "Amphitremida (poche, 1913) is a new major, ubiquitous labyrinthulomycete clade". PLoS One. 8 (1): e53046. Bibcode:2013PLoSO...853046G. doi:10.1371/journal.pone.0053046. PMC 3544814. PMID 23341921. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3544814" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3544814</a> <a href="#fnref:11" class="footnote-back-ref">↩</a></p></li> <li id="fn:12"><p>Anderson, O. Roger; Cavalier-Smith, Thomas (2012). "Ultrastructure of Diplophrys parva, a New Small Freshwater Species, and a Revised Analysis of Labyrinthulea (Heterokonta)". Acta Protozoologica. 8 (1): 291–304. doi:10.4467/16890027AP.12.023.0783. <a href="http://www.eko.uj.edu.pl/ap" target="_blank">http://www.eko.uj.edu.pl/ap</a> <a href="#fnref:12" class="footnote-back-ref">↩</a></p></li> <li id="fn:13"><p>FioRito, Rebecca; Leander, Celeste; Leander, Brian (2016). "Characterization of three novel species of Labyrinthulomycota isolated from ochre sea stars (Pisaster ochraceus)". Marine Biology. 163 (8): 170. doi:10.1007/s00227-016-2944-5. S2CID 43399688. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:13" class="footnote-back-ref">↩</a></p></li> <li id="fn:14"><p>Adl SM, Bass D, Lane CE, Lukeš J, Schoch CL, Smirnov A, Agatha S, Berney C, Brown MW, Burki F, Cárdenas P, Čepička I, Chistyakova L, del Campo J, Dunthorn M, Edvardsen B, Eglit Y, Guillou L, Hampl V, Heiss AA, Hoppenrath M, James TY, Karnkowska A, Karpov S, Kim E, Kolisko M, Kudryavtsev A, Lahr DJG, Lara E, Le Gall L, Lynn DH, Mann DG, Massana R, Mitchell EAD, Morrow C, Park JS, Pawlowski JW, Powell MJ, Richter DJ, Rueckert S, Shadwick L, Shimano S, Spiegel FW, Torruella G, Youssef N, Zlatogursky V, Zhang Q (2019). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. doi:10.1111/jeu.12691. PMC 6492006. PMID 30257078. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6492006" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6492006</a> <a href="#fnref:14" class="footnote-back-ref">↩</a></p></li> <li id="fn:15"><p>Bennett, Reuel M.; Honda, D.; Beakes, Gordon W.; Thines, Marco (2017). "Chapter 14. Labyrinthulomycota". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Springer. pp. 507–542. doi:10.1007/978-3-319-28149-0_25. ISBN 978-3-319-28147-6. <a href="978-3-319-28147-6" target="_blank">978-3-319-28147-6</a> <a href="#fnref:15" class="footnote-back-ref">↩</a></p></li> <li id="fn:16"><p>Bennett, Reuel M.; Honda, D.; Beakes, Gordon W.; Thines, Marco (2017). "Chapter 14. Labyrinthulomycota". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. 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