Plasmodiophore

The plasmodiophores[1] (also known as plasmophorids[2] or plasmodiophorids[3]) are a group of obligate endoparasitic protists belonging to the subphylum Endomyxa in Cercozoa.[4] Taxonomically, they are united under a single family Plasmodiophoridae, order Plasmodiophorida, sister to the phagomyxids.[5]

Plasmodiophores
"Plasmodiophoraceae" 1960 illustration.
Scientific classification Edit this classification
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Clade: Rhizaria
Phylum: Endomyxa
Class: Phytomyxea
Order: Plasmodiophorida
Cook, 1928
Family: Plasmodiophoridae
Zopf, 1884
Type genus
Plasmodiophora
M. Woronin, 1877
Genera
Synonyms
  • Plasmodiophoromycota Whittaker, 1969
  • Plasmodiophoromycetes K. Cejp, 1957
  • Plasmodiophorales Engl., 1892
  • Plasmodiophorales F.L. Stevens, 1913
  • Woroninales A.A. Jaczewski & P.A. Jaczewski, 1931
  • Plasmodiophoraceae Zopf ex Berl., 1888
  • Woroninaceae M. von Minden, 1911

Ecology and pathology

Plasmodiophores are pathogenic for a wide range of organisms, but mainly green plants. The more commonly recognized are agents of plant diseases such as clubroot, powdery scab and crook root of watercress,[6] or vectors for viruses that infect beets, peanut, monocots and potatoes, such as the potato mop-top virus or the beet necrotic yellow vein virus.[7][8]

Taxonomy

History

The plasmodiophores have historically been regarded as Fungi. The first description of plasmodiophores as a taxonomic group was in 1885 by Zopf, who united two genera Plasmodiophora and Tetramyxa in a common family “Plasmodiophoreæ”, inside the group “Monadineæ”, as part of the division Myxomycetes. The family was renamed “Plasmodiophoraceae” in 1888 by Berlese.[9] In 1892, Engler placed the family in its own class “Plasmodiophorales”, later renamed “Plasmodiophoromycetes” to fit nomenclature standards.[10]

In 1969 Whittaker, in his five-kingdom system, elevated the group to a separate phylum “Plasmodiophoromycota”, acknowledging them as protists instead of fungi.[1]

In 1993 Cavalier-Smith included the plasmodiophores and their sister group Phagomyxida in their current class, Phytomyxea, as part of a polyphyletic phylum called Opalozoa, which at the time contained a diverse assemblage of unrelated zooflagellates, opalines and proteomyxids.[11] Eventually this phylum was discarded, and the name Opalozoa was modified to label a group inside the phylum Bigyra containing the opalines, bicosoecids and related organisms.[12]

Finally, after phylogenetic analyses, in 2002 Cavalier-Smith placed all Phytomyxea, including plasmodiophores, in the subphylum Endomyxa, inside the rhizarian phylum Cercozoa.[13][14]

Classification

The number of genera varies between sources. There are three accepted genera in the group according to the WoRMS register: Plasmodiophora, Spongospora and Tetramyxa.[15] Below is a complete list with genera that are not included in the register but appear in relevant sources:[3][16][17]

  • Ligniera R. Maire & A. Tison 1911 (=Anisomyxa Nemec 1913; Rhizomyxa Borzí 1884; Sorolpidium B. Nĕmec 1911)
  • Membranosorus C.H. Ostenfeld & H.E. Petersen 1930
  • Octomyxa J.N. Couch J. Leitner & A. Whiffen 1939
  • Polymyxa G.A. Ledingham 1939
  • Plasmodiophora Woronin 1877 (=Ostenfeldiella Ferdinandsen & Winge 1914)
  • Sorodiscus G. Lagerheim & Ø. Winge 1912
  • Sorosphaerula J. Schröt. 1886 (as Sorosphaera) nom. nov. Neuhauser & Kirchmair 2011[18] (=Tuburcinia E.M. Fries; Sorosporium F. Rudolphi 1829)
  • Spongospora Brunch. 1887 (=Clathrosorus C. Ferdinandson & Ö. Winge 1920)
  • Tetramyxa K. Goebel 1884 (=Molliardia R. Maire & A. Tison 1911; Thecaphora W. A. Setchell 1924)
  • Woronina Cornu 1872

These genera were once considered plasmodiophores[19] until they were excluded:[17]

  • Cystospora J.E. Elliott 1916 – possibly a physiological symptom.
  • Frankiella Maire & A. Tison 1909 – synonym of the bacteria Frankia.
  • Peltomyces L. Léger 1909 – excluded as unclassifiable.
  • Pyrrhosorus H. O. Juel 1901 – considered Labyrinthulida incertae sedis.[20]
  • Sporomyxa L. Léger 1907 – excluded as unclassifiable.
  • Trematophlyctis Patouillard 1918 – a chytrid fungus.[21]

References

  1. Whittaker RH (10 January 1969). "New concepts of kingdoms or organisms. Evolutionary relations are better represented by new classifications than by the traditional two kingdoms". Science. 163 (3863): 150–60. doi:10.1126/science.163.3863.150. PMID 5762760.
  2. Stjelja S, Fogelqvist J, Tellgren-Roth C, et al. (2019). "The architecture of the Plasmodiophora brassicae nuclear and mitochondrial genomes". Sci Rep. 9 (15753): 15753. doi:10.1038/s41598-019-52274-7. PMC 6823432. PMID 31673019.
  3. Neuhauser S, Kirchmair M, Bulman S, et al. (23 February 2014). "Cross-kingdom host shifts of phytomyxid parasites". BMC Evol Biol. 14 (33): 33. doi:10.1186/1471-2148-14-33. PMC 4016497. PMID 24559266.
  4. Irwin, Nicholas A.T.; Tikhonenkov, Denis V.; Hehenberger, Elisabeth; Mylnikov, Alexander P.; Burki, Fabien; Keeling, Patrick J. (2019-01-01). "Phylogenomics supports the monophyly of the Cercozoa". Molecular Phylogenetics and Evolution. 130: 416–423. doi:10.1016/j.ympev.2018.09.004. ISSN 1055-7903. PMID 30318266. S2CID 52982396.
  5. Cavalier-Smith, Thomas; Chao, Ema E.; Lewis, Rhodri (April 2018). "Multigene phylogeny and cell evolution of chromist infrakingdom Rhizaria: contrasting cell organisation of sister phyla Cercozoa and Retaria". Protoplasma. 255 (5): 1517–1574. doi:10.1007/s00709-018-1241-1. PMC 6133090. PMID 29666938.
  6. Braselton JP (2001). "Plasmodiophoromycota". In McLaughlin DJ, McLaughlin EG, Lemke PA (eds.). The Mycota. Vol. VII: Systematics and Evolution Part A. Springer, Berlin, Heidelberg. pp. 81–91. doi:10.1007/978-3-662-10376-0_4. ISBN 978-3-662-10376-0.
  7. Schwelm A, Badstöber J, Bulman S, Desoignies N, Etemadi M, Falloon RE, Gachon CMM, Legreve A, Lukeš J, Merz U, Nenarokova A, Strittmatter M, Sullivan BK, Neuhauser S (April 2018). "Not in your usual Top 10: protists that infect plants and algae". Mol Plant Pathol. 19 (4): 1029–1044. doi:10.1111/mpp.12580. PMC 5772912. PMID 29024322.
  8. Neuhauser, Sigrid; Kirchmair, Martin; Gleason, Frank H. (28 April 2011). "Ecological roles of the parasitic phytomyxids (plasmodiophorids) in marine ecosystems – a review". Marine and Freshwater Research. 62 (4): 365–371. doi:10.1071/MF10282. PMC 3272469. PMID 22319023.
  9. Saccardo PA, Traverso GB, Trotter A (1882). Sylloge fungorum omnium hucusque cognitorum. Vol. 1. p. 323. doi:10.5962/bhl.title.5371. LCCN agr12002244. OCLC 2472326.
  10. Engler A (1903). Syllabus der Pflanzenfamilien : eine Übersicht über das gesamte Pflanzensystem mit Berücksichtigung der Medicinal- und Nutzpflanzen nebst einer Übersicht über die Florenreiche und Florengebiete der Erde zum Gebrauch bei Vorlesungen und Studien über specielle und medicinisch-pharmaceutische Botanik (3rd ed.). Berlin: Borntraeger. doi:10.5962/bhl.title.22956.
  11. Cavalier-Smith T (September 1993). "The Protozoan Phylum Opalozoa". Eukaryotic Microbiology. 40 (5): 609–615. doi:10.1111/j.1550-7408.1993.tb06117.x. S2CID 84129692.
  12. Cavalier-Smith, Thomas; Scoble, Josephine Margaret (2012). "Phylogeny of Heterokonta: Incisomonas marina, a uniciliate gliding opalozoan related to Solenicola (Nanomonadea), and evidence that Actinophryida evolved from raphidophytes". European Journal of Protistology. 49 (3): 328–353. doi:10.1016/j.ejop.2012.09.002. PMID 23219323.
  13. Cavalier-Smith T (1 March 2002). "The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa". International Journal of Systematic and Evolutionary Microbiology. 52 (2): 297–354. doi:10.1099/00207713-52-2-297. PMID 11931142.
  14. Cavalier-Smith E, Chao EE (2003). "Phylogeny and Classification of Phylum Cercozoa (Protozoa)". Protist. 154 (3–4): 341–358. doi:10.1078/143446103322454112. ISSN 1434-4610. PMID 14658494.
  15. "Plasmodiophoridae". WoRMS. World Register of Marine Species. Retrieved 19 October 2022.
  16. Sparrow FK (1960). Aquatic Phycomycetes. University of Michigan studies, Scientific series, v. 15. Ann Arbor, University of Michigan Press. doi:10.5962/bhl.title.5685. LCCN 59011269. OCLC 1362167.
  17. Dick, Michael W. (2001). Straminipilous Fungi: Systematics of the Peronosporomycetes Including Accounts of the Marine Straminipilous Protists, the Plasmodiophorids and Similar Organisms (1 ed.). Springer Dordrecht. doi:10.1007/978-94-015-9733-3. ISBN 978-94-015-9733-3. S2CID 28755980.
  18. Neuhauser, Sigrid; Kirchmair, Martin (18 August 2011). "Sorosphaerula nom. n. for the Plasmodiophorid Genus Sorosphaera J. Schröter 1886 (Rhizaria: Endomyxa: Phytomyxea: Plasmodiophorida)". Journal of Eukaryotic Microbiology. 58 (5): 469–470. doi:10.1111/j.1550-7408.2011.00571.x. PMC 3245851. PMID 21851471.
  19. "Plasmodiophoraceae". www.mycobank.org.
  20. Juel. "Pyrrhosorus". WoRMS. World Register of Marine Species.
  21. Wijayawardene, N.N.; Hyde, K.D.; Dai, D.Q.; Sánchez-García, M.; Goto, B.T.; Saxena, R.K.; et al. (2022). "Outline of Fungi and fungus-like taxa – 2021". Mycosphere. 13 (1): 53–453. doi:10.5943/mycosphere/13/1/2. S2CID 249054641.
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