Plasmodiophores | |
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"Plasmodiophoraceae" 1960 illustration. | |
Scientific 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 | |
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 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]
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]
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]
These genera were once considered plasmodiophores [19] until they were excluded: [17]
Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead united by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eukaryotic group to be recognized mainly through molecular phylogenies. They are the natural predators of many species of bacteria. They are closely related to the phylum Retaria, comprising amoeboids that usually have complex shells, and together form a supergroup called Rhizaria.
The Rhizaria are a diverse and species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has been described. This group was used by Cavalier-Smith in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon. Being described mainly from rDNA sequences, they vary considerably in form, having no clear morphological distinctive characters (synapomorphies), but for the most part they are amoeboids with filose, reticulose, or microtubule-supported pseudopods. In the absence of an apomorphy, the group is ill-defined, and its composition has been very fluid. Some Rhizaria possess mineral exoskeletons, which are in different clades within Rhizaria made out of opal, celestite, or calcite. Certain species can attain sizes of more than a centimeter with some species being able to form cylindrical colonies approximately 1 cm in diameter and greater than 1 m in length. They feed by capturing and engulfing prey with the extensions of their pseudopodia; forms that are symbiotic with unicellular algae contribute significantly to the total primary production of the ocean.
Phaeodarea or Phaeodaria is a group of amoeboid cercozoan organisms. They are traditionally considered radiolarians, but in molecular trees do not appear to be close relatives of the other groups, and are instead placed among the Cercozoa. They are distinguished by the structure of their central capsule and by the presence of a phaeodium, an aggregate of waste particles within the cell.
The Phytomyxea are a class of parasites that are cosmopolitan, obligate biotrophic protist parasites of plants, diatoms, oomycetes and brown algae. They are divided into the orders Plasmodiophorida and Phagomyxida. Plasmodiophorids are best known as pathogens or vectors for viruses of arable crops.
Monadofilosa is a grouping of Cercozoa. These organisms are single-celled amoeboid protists.
Endomyxa is a group of eukaryotic organisms in the supergroup Rhizaria. They were initially a subphylum of Cercozoa and later a subphylum of Retaria, but several analyses have proven they are a phylogenetically separate lineage, and Endomyxa is currently regarded as its own phylum.
Thecofilosea is a class of unicellular testate amoebae belonging to the phylum Cercozoa. They are amoeboflagellates, organisms with flagella and pseudopodia, distinguished from other cercozoa by their scale-lacking test composed of organic material. They are closely related to the Imbricatea, a group of testate amoebae with tests composed of inorganic silica scales.
Bigyra is a phylum of microscopic eukaryotes that are found at the base of the Stramenopiles clade. It includes three well-known heterotrophic groups Bicosoecida, Opalinata and Labyrinthulomycetes, as well as several small clades initially discovered through environmental DNA samples: Nanomonadea, Placididea, Opalomonadea and Eogyrea. The classification of Bigyra has changed several times since its origin, and its monophyly remains unresolved.
The sarcomonads or class Sarcomonadea are a group of amoeboid biciliate protists in the phylum Cercozoa. They are characterized by a propensity to move through gliding on their posterior cilium or through filopodia, a lack of scales or external theca, a soft cell surface without obvious cortical filamentous or membranous skeleton, two cilia without scales or hairs, tubular mitochondrial cristae, near-spherical extrusomes, and a microbody attached to the nucleus.
Plasmodiophora is a genus in class Phytomyxea.
The vampyrellids, colloquially known as vampire amoebae, are a group of free-living predatory amoebae classified as part of the lineage Endomyxa. They are distinguished from other groups of amoebae by their irregular cell shape with propensity to fuse and split like plasmodial organisms, and their life cycle with a digestive cyst stage that digests the gathered food. They appear worldwide in marine, brackish, freshwater and soil habitats. They are important predators of an enormous variety of microscopic organisms, from algae to fungi and animals. They are also known as aconchulinid amoebae.
The initial version of a classification system of life by British zoologist Thomas Cavalier-Smith appeared in 1978. This initial system continued to be modified in subsequent versions that were published until he died in 2021. As with classifications of others, such as Carl Linnaeus, Ernst Haeckel, Robert Whittaker, and Carl Woese, Cavalier-Smith's classification attempts to incorporate the latest developments in taxonomy., Cavalier-Smith used his classifications to convey his opinions about the evolutionary relationships among various organisms, principally microbial. His classifications complemented his ideas communicated in scientific publications, talks, and diagrams. Different iterations might have a wider or narrow scope, include different groupings, provide greater or lesser detail, and place groups in different arrangements as his thinking changed. His classifications has been a major influence in the modern taxonomy, particularly of protists.
Leucodictyids are heterotrophic amoeboid protists that comprise the order Leucodictyida in the phylum Cercozoa.
Ligniera is a protist genus of the family Plasmodiophoraceae.
Maullinia is a genus of intracellular, phytomyxid parasites found across the Southern Hemisphere though primarily in Chile, The Prince Edward Islands, South Africa, Australia, and New Zealand. These parasites infiltrate the cells of their brown algal hosts via cytoplasmic extensions called plasmodia that divide synchronously, becoming increasingly multi-nucleate and engulfing the host cell organelles as they grow. Eventually, as the plasmodia fill the entire cell volume, the host cells become hypertrophied and grow to 3- 4x their original size, showing up as swollen appendages or galls on the host tissue at a macroscopic level. These swollen regions will burst alongside the mature Maullinia plasmodia, releasing biflagellated zoospores to the inter- and extracellular space to disperse the infection further. Zoospores can come from sporangial plasmodia, as in M. ectocarpii, or from resting spores, as in M. braseltonii.
Phagomyxids are a group of obligate endoparasitic protists belonging to the subphylum Endomyxa in Cercozoa. Taxonomically, they are united under a single family Phagomyxidae, order Phagomyxida, sister to the plasmodiophores.
Tetramyxa is a cercozoan protist, member of the plasmodiophores, parasite of several flowering plants. It was first described by Karl von Goebel in 1884, in his work Flora. The genus is characterized by the appearance of resting spores in groups of four.
Tetramyxa parasitica is a species of parasitic cercozoan, member of the plasmodiophorids, that causes gall formation on multiple genera of aquatic plants. It was first discovered on roots of Ruppia and described by Karl von Goebel in 1884 in his work Flora, where it became the type species of the genus Tetramyxa.
Massisteriidae is a family of granofilosean protists within the phylum Cercozoa.
Euglyphia is a group of imbricate protists in the phylum Cercozoa. They are unicellular eukaryotes characterized by a cell body covered in large imbricate scales, and an apical aperture through which they extend either filose pseudopodia or two cilia of different sizes that are not used for gliding.