Stramenopile

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Stramenopiles
Temporal range: Late Mesoproterozoic-present, 1025–0 Ma [1]
Stramenopiles diversity.png
Diversity of stramenopiles
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Clade: Stramenopiles
Patterson 1989 [2] emend. Adl et al. 2005 [3]
Phyla and subphyla [4]
Diversity
>100000 species [1]
Synonyms

The stramenopiles, also called heterokonts, are a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have been secondarily lost (in which case relatedness to stramenopile ancestors is evident from other shared cytological features or from genetic similarity). Stramenopiles represent one of the three major clades in the SAR supergroup, along with Alveolata and Rhizaria.

Contents

Stramenopiles are eukaryotes; most are single-celled, but some are multicellular including some large seaweeds, the brown algae. The group includes a variety of algal protists, heterotrophic flagellates, opalines and closely related proteromonad flagellates (all endobionts in other organisms); the actinophryid Heliozoa, and oomycetes. The tripartite hairs characteristic of the group have been lost in some of the included taxa – for example in most diatoms.

Many stramenopiles are unicellular flagellates, and most others produce flagellated cells at some point in their lifecycles, for instance as gametes or zoospores. Most flagellated heterokonts have two flagella; the anterior flagellum has one or two rows of stiff hairs or mastigonemes, and the posterior flagellum is without such embellishments, being smooth, usually shorter, or in a few cases not projecting from the cell.

The term 'heterokont' is used both as an adjective – indicating that a cell has two dissimilar flagella, and as the name of a taxon. The groups included in that taxon have however varied widely, creating the 'heterokont problem', now resolved by the definition of the stramenopiles.

History

The term 'stramenopile' was introduced by D. J. Patterson in 1989, defining a group that overlapped with the ambiguously defined heterokonts. [12] [13] The name "stramenopile" has been discussed by J. C. David. [14]

The heterokont problem

The term 'heterokont' is used as both an adjective – indicating that a cell has two dissimilar flagella – and as the name of a taxon. The taxon 'Heterokontae' was introduced in 1899 by Alexander Luther for algae that are now considered the Xanthophyceae. [15] But the same term was used for other groupings of algae. For example, in 1956, Copeland [16] used it to include the xanthophytes (using the name Vaucheriacea), a group that included what became known as the chrysophytes, the silicoflagellates, and the hyphochytrids. Copeland also included the unrelated collar flagellates (as the choanoflagellates) in which he placed the bicosoecids. He also included the not-closely related haptophytes. The consequence of associating multiple concepts to the taxon 'heterokont' is that the meaning of 'heterokont' can only be made clear by making reference to its usage: Heterokontae sensu Luther 1899; Heterokontae sensu Copeland 1956, etc. This contextual clarification is rare, such that when the taxon name is used, it is unclear how it should be understood. The term 'heterokont' has lost its usefulness in critical discussions about the identity, nature, character and relatedness of the group. [17] The term 'stramenopile' sought to identify a clade (monophyletic and holophyletic lineage) using the approach developed by transformed cladists of pointing to a defining innovative characteristic or apomorphy. [18]

Over time, the scope of application has changed, especially when in the 1970s ultrastructural studies revealed greater diversity among the algae with chromoplasts (chlorophylls a and c) than had previously been recognized. At the same time, a protistological perspective was replacing the 19th century one based on the division of unicellular eukaryotes into animals and plants. One consequence was that an array of heterotrophic organisms, many not previously considered as 'heterokonts', were seen as related to the 'core heterokonts' (those having anterior flagella with stiff hairs). Newly recognized relatives included the parasitic opalines, proteromonads, and actinophryid Heliozoa. They joined other heterotrophic protists, such as bicosoecids, labyrinthulids, and oomycete fungi, that were included by some as heterokonts and excluded by others. Rather than continue to use a name whose meaning had changed over time and was hence ambiguous, the name 'stramenopile' was introduced to refer to the clade of protists that had tripartite stiff (usually flagellar) hairs and all their descendants. Molecular studies confirm that the genes that code for the proteins of these hairs are exclusive to stramenopiles. [19]

Characteristics

Cafeteria roenbergensis FENCHEL and D J PATTERSON schematic drawing.svg
Schematic drawing of Cafeteria roenbergensis (a heterotrophic bicosoecid), a common bacterivore in marine ecosystems: the anterior flagellum is tripartite and covered with hairs (mastigonemes); the posterior flagellum is without hairs.
Cafeteria roenbergensis atcc50561 Protsville.jpg
Two living C. roenbergensis. Light micrograph. The cells are about 6 µm long. The anterior flagellum beats with an undulating pattern, the posterior (recurrent or smooth) flagellum usually holds the cell to the substrate.

The presumed apomorphy of tripartite flagellar hairs in stramenopiles is well characterized. The basal part of the hair is flexible and inserts into the cell membrane; the second part is dominated by a long stiff tube (the 'straw' or 'stramen'); and finally the tube is tipped by many delicate hairs called mastigonemes. [20] The proteins that code for the mastigonemes appear to be exclusive to the stramenopile clade, and are present even in taxa (such as diatoms) that no longer have such hairs. [21]

Most stramenopiles have two flagella near the apex. [22] They are usually supported by four microtubule roots in a distinctive pattern. There is a transitional helix inside the flagellum where the beating axoneme with its distinctive geometric pattern of nine peripheral couplets around two central microtubules changes into the nine-triplet structure of the basal body. [23]

Plastids

Many stramenopiles have plastids which enable them to photosynthesise, using light to make their own food. Those plastids are coloured off-green, orange, golden or brown because of the presence of chlorophyll a, chlorophyll c, and fucoxanthin. This form of plastid is called a stramenochrome or chromoplast. [a] The most significant autotrophic stramenopiles are the brown algae (wracks and many other seaweeds), and the diatoms. The latter are among the most significant primary producers in marine and freshwater ecosystems. [24] Most molecular analyses suggest that the most basal stramenopiles lacked plastids and were accordingly colourless heterotrophs, feeding on other organisms. This implies that the stramenopiles arose as heterotrophs, diversified, and then some of them acquired chromoplasts. Some lineages (such as the axodine lineage that included the chromophytic pedinellids, colourless ciliophryids, and colourless actinophryid heliozoa) have secondarily reverted to heterotrophy. [25] [26]

Ecology

Giant kelp, Macrocystis pyrifera, an example of a multicellular stramenopile, is a large seaweed, up to 45 metres (150 feet) long, in the Phaeophyceae, within the Gyrista. Kelp-forest-Monterey.jpg
Giant kelp, Macrocystis pyrifera , an example of a multicellular stramenopile, is a large seaweed, up to 45 metres (150 feet) long, in the Phaeophyceae, within the Gyrista.

Some stramenopiles are significant as autotrophs and as heterotrophs in natural ecosystems; others are parasitic. Blastocystis is a gastrointestinal parasite of humans; [27] opalines and proteromonads live in the intestines of cold-blooded vertebrates and have been described as parasitic; [28] oomycetes include some significant plant pathogens such as the cause of potato blight, Phytophthora infestans . [29] Diatoms are major contributors to global carbon cycles because they are the most important autotrophs in most marine habitats. [30] The brown algae, including familiar seaweeds like wrack and kelp, are major autotrophs of the intertidal and subtidal marine habitats. [31] Some of the bacterivorous stramenopiles, such as Cafeteria, are common and widespread consumers of bacteria, and thus play a major role in recycling carbon and nutrients within microbial food webs. [32] [33]

Evolution

External

Stramenopiles are most closely related to alveolates and Rhizaria, all of which have tubular mitochondrial cristae and collectively form the SAR supergroup, whose name is formed from their initials. [34] [26] [35] The ancestor of the SAR supergroup appears to have captured a unicellular photosynthetic red alga, and many stramenopiles, as well as members of other SAR groups such as the Rhizaria, still have plastids which retain the double membrane of the red alga and a double membrane surrounding it, for a total of four membranes. [36] In addition, species of Telonemia, the sister group to SAR, exhibit heterokont flagella with tripartite mastigonemes, implying a more ancient origin of stramenopile characteristics. [37]

TSAR

Internal

The following cladogram summarizes the evolutionary relationships between stramenopiles. The phylogenetic relationships of Bigyra vary greatly from one analysis to the next: it has been recovered as either monophyletic [38] [39] or paraphyletic. When paraphyletic, the branching order of the bigyran groups also varies: in some studies, Sagenista is the most basal-branching clade, [38] [40] [41] while in others Opalozoa is the most basal. [42] Nonetheless, Platysulcea is consistently recovered as the sister clade to all other stramenopiles. [39] [40] In addition, a flagellate species discovered in 2023, Kaonashia insperata , remains in an uncertain phylogenetic position, but more closely related to Gyrista than to other clades. [41]

Classification

Electron micrograph of the protist Paraphysomonas butcheri. It illustrates the stramenopile property - of having stiff hairs. The hairs attach to one longer flagellum, the other is without hairs (an arrangement also called 'heterokont', meaning "unequal"). The body of the flagellate is coated with delicate scales. Paraphysomonas feeds on bacteria, two of which lie near the hairy flagellum. Paraphysomonas butcheri whole mount.jpg
Electron micrograph of the protist Paraphysomonas butcheri. It illustrates the stramenopile property – of having stiff hairs. The hairs attach to one longer flagellum, the other is without hairs (an arrangement also called 'heterokont', meaning "unequal"). The body of the flagellate is coated with delicate scales. Paraphysomonas feeds on bacteria, two of which lie near the hairy flagellum.

The classification of the stramenopiles according to Adl et al. (2019), with additions from newer research: [43] [4]

Notes

  1. ↑ They are not called chloroplasts, the most common form of photosynthetic plastid. If used narrowly, a chloroplast is a plastid which contains chlorophyll B, as in green algae, some euglenids, and the land plants.

Related Research Articles

<span class="mw-page-title-main">Flagellate</span> Group of protists with at least one whip-like appendage

A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word flagellate also describes a particular construction characteristic of many prokaryotes and eukaryotes and their means of motion. The term presently does not imply any specific relationship or classification of the organisms that possess flagella. However, several derivations of the term "flagellate" are more formally characterized.

<span class="mw-page-title-main">Alveolate</span> Superphylum of protists

The alveolates are a group of protists, considered a major clade and superphylum within Eukarya. They are currently grouped with the stramenopiles and Rhizaria among the protists with tubulocristate mitochondria into the SAR supergroup.

<span class="mw-page-title-main">Cryptomonad</span> Group of algae and colorless flagellates

The cryptomonads are a group of algae, most of which have plastids. They are traditionally considered a division of algae among phycologists, under the name of Cryptophyta. They are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, with an anterior groove or pocket. At the edge of the pocket there are typically two slightly unequal flagella. Some may exhibit mixotrophy. They are classified as clade Cryptomonada, which is divided into two classes: heterotrophic Goniomonadea and phototrophic Cryptophyceae. The two groups are united under three shared morphological characteristics: presence of a periplast, ejectisomes with secondary scroll, and mitochondrial cristae with flat tubules. Genetic studies as early as 1994 also supported the hypothesis that Goniomonas was sister to Cryptophyceae. A study in 2018 found strong evidence that the common ancestor of Cryptomonada was an autotrophic protist.

<span class="mw-page-title-main">Axodine</span> Class of single-celled organisms

The axodines are a group of unicellular stramenopiles that includes silicoflagellate and rhizochromulinid algae, actinomonad heterotrophic flagellates and actinophryid heliozoa. Alternative classifications treat the dictyochophytes as heterokont algae, or as Chrysophyceae. Other overlapping taxonomic concepts include the Actinochrysophyceae, Actinochrysea or Dictyochophyceae sensu lato. The grouping was proposed on the basis of ultrastructural similarities, and is consistent with subsequent molecular comparisons.

<span class="mw-page-title-main">Chromista</span> Eukaryotic biological kingdom

Chromista is a proposed but polyphyletic biological kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles (plastids). It includes all eukaryotes whose plastids contain chlorophyll c and are surrounded by four membranes. If the ancestor already possessed chloroplasts derived by endosymbiosis from red algae, all non-photosynthetic Chromista have secondarily lost the ability to photosynthesise. Its members might have arisen independently as separate evolutionary groups from the last eukaryotic common ancestor.

<span class="mw-page-title-main">Labyrinthulomycetes</span> Class of protists that produce a filamentous network

Labyrinthulomycetes (ICBN) or Labyrinthulea (ICZN) is a class of protists that produce a network of filaments or tubes, which serve as tracks for the cells to glide along and absorb nutrients for them. The two main groups are the labyrinthulids and thraustochytrids. They are mostly marine, commonly found as parasites on algae and seagrasses or as decomposers on dead plant material. They also include some parasites of marine invertebrates and mixotrophic species that live in a symbiotic relationship with zoochlorella.

<span class="mw-page-title-main">Telonemia</span> Phylum of single-celled organisms

Telonemia is a phylum of microscopic eukaryotes commonly known as telonemids. They are unicellular free-living flagellates with a unique combination of cell structures, including a highly complex cytoskeleton unseen in other eukaryotes.

<span class="mw-page-title-main">Ochrophyte</span> Phylum of algae

Ochrophytes, also known as heterokontophytes or stramenochromes, are a group of algae. They are the photosynthetic stramenopiles, a group of eukaryotes, organisms with a cell nucleus, characterized by the presence of two unequal flagella, one of which has tripartite hairs called mastigonemes. In particular, they are characterized by photosynthetic organelles or plastids enclosed by four membranes, with membrane-bound compartments called thylakoids organized in piles of three, chlorophyll a and c as their photosynthetic pigments, and additional pigments such as β-carotene and xanthophylls. Ochrophytes are one of the most diverse lineages of eukaryotes, containing ecologically important algae such as brown algae and diatoms. They are classified either as phylum Ochrophyta or Heterokontophyta, or as subphylum Ochrophytina within phylum Gyrista. Their plastids are of red algal origin.

<span class="mw-page-title-main">Protist</span> Eukaryotes other than animals, plants or fungi

A protist or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a polyphyletic grouping of several independent clades that evolved from the last eukaryotic common ancestor.

<span class="mw-page-title-main">SAR supergroup</span> Eukaryotes superphylum

SAR or Harosa is a highly diverse clade of eukaryotes, often considered a supergroup, that includes stramenopiles (heterokonts), alveolates, and rhizarians. It is a node-based taxon, including all descendants of the three groups' last common ancestor, and comprises most of the now-rejected Chromalveolata. Their sister group has been found to be telonemids, with which they make up the TSAR clade.

<span class="mw-page-title-main">Bigyra</span> Phylum of single-celled organisms

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.

<span class="mw-page-title-main">Katablepharid</span> Group of algae

The kathablepharids or katablepharids are a group of heterotrophic flagellates closely related to cryptomonads. First described by Heinrich Leonhards Skuja in 1939, kathablepharids were named after the genus Kathablepharis. This genus is corrected to Katablepharis under botanical nomenclature, but the original spelling is maintained under zoological nomenclature. They are single-celled protists with two anteriorly directed flagella, an anterior cytostome for ingesting eukaryotic prey, and a sheath that covers the cell membrane. They have extrusomes known as ejectisomes, as well as tubular mitochondrial cristae.

<span class="mw-page-title-main">Haptista</span> Group of protists

Haptista is a proposed group of protists made up of centrohelids and haptophytes. Phylogenomic studies indicate that Haptista, together with Ancoracysta twista, forms a sister clade to the SAR+Telonemia supergroup, but it may also be sister to the Cryptista (+Archaeplastida). It is thus one of the earliest diverging Diaphoretickes.

Platysulcus tardus is an eukaryotic microorganism that was recently discovered to be the earliest diverging lineage of the Heterokont phylogenetic tree. It is the only member of the family Platysulcidae, order Platysulcida and class Platysulcea.

Endohelea is a proposed clade of eukaryotes that are related to Archaeplastida and the SAR supergroup. They used to be considered heliozoans, but phylogenetically they belong to a group of microorganisms known as Cryptista.

<span class="mw-page-title-main">Ultrastructural identity</span>

Ultrastructural identity is a concept in biology. It asserts that evolutionary lineages of eukaryotes in general and protists in particular can be distinguished by complements and arrangements of cellular organelles. These ultrastructural components can be visualized by electron microscopy.

Thecamonadinae is a subfamily of heterotrophic protists. It is a monophyletic group, or clade, of apusomonads, a group of protozoa with two flagella closely related to the eukaryotic supergroup Opisthokonta. The subfamily contains two genera Chelonemonas and Thecamonas, which are found in marine habitats.

Commation is a genus of marine heterotrophic protists closely related to the actinophryids. It contains two species, Commation cryoporinum and Commation eposianum, discovered in antarctic waters and described in 1993. Currently, the genus is classified within a monotypic family Commatiidae and order Commatiida. Along with the photosynthetic raphidophytes, these organisms compose the class of stramenopiles known as Raphidomonadea.

<span class="mw-page-title-main">Colponemid</span> Group of predatorial flagellates

Colponemids are free-living alveolates, unicellular flagellates related to dinoflagellates, apicomplexans and ciliates. They are predators of other small eukaryotes, found in freshwater, marine and soil environments. They do not form a solid clade, but a sparse group of deep-branching alveolate lineages.

<span class="mw-page-title-main">Anisonemia</span> Group of flagellates

Anisonemia is a clade of single-celled protists belonging to the phylum Euglenozoa, relatives of the Euglenophyceae algae. They are flagellates, with two flagella for locomotion. Anisonemia includes various phagotrophic species and a group of primary osmotrophic protists known as Aphagea.

References

  1. 1 2 H.S. Yoon; R.A. Andersen; S.M. Boo; D. Bhattacharya (17 February 2009). "Stramnenopiles". Encyclopedia of Microbiology (Third Edition): 721–731. doi:10.1016/B978-012373944-5.00253-4 . Retrieved 2 March 2024.
  2. ↑ Patterson, D.J. (1989). "Stramenopiles: Chromophytes from a protistan perspective". In Green, J.C.; Leadbeater, B.S.C.; Diver, W.L. (eds.). The chromophyte algae: Problems and perspectives . Clarendon Press. ISBN   978-0198577133.
  3. ↑ Sina M Adl; Alastair G B Simpson; Mark A Farmer; et al. (1 September 2005). "The new higher level classification of eukaryotes with emphasis on the taxonomy of protists". Journal of Eukaryotic Microbiology . 52 (5): 399–451. doi:10.1111/J.1550-7408.2005.00053.X. ISSN   1066-5234. PMID   16248873. Wikidata   Q22065654.
  4. 1 2 Cavalier-Smith, Thomas (January 2018). "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.
  5. ↑ Vørs, N. (1993). "Marine heterotrophic amoebae, flagellates and heliozoa from Belize (Central America) and Tenerife". Journal of Eukaryotic Microbiology. 40 (3): 272–287. doi:10.1111/j.1550-7408.1993.tb04917.x. S2CID   221852241.
  6. ↑ David, J.C. (2002). "A preliminary catalogue of the names of fungi above the rank of order". Constancea. 83: 1–30.
  7. ↑ Cavalier-Smith, T. (1999). "The kingdom Chromista, origin and systematics". In Round, F.E.; Chapman, D.J. (eds.). Progress in Phycological Research. Vol. 4. Elsevier. pp. 309–347. ISBN   978-0-948737-00-8.
  8. ↑ van den Hoek, C.; Mann, D.G.; Jahns, H.M. (1995). Algae An Introduction to Phycology. Cambridge University Press. ISBN   978-0-521-30419-1.
  9. ↑ Alexopoulos, C.J.; Mims, C.W.; Blackwell, M. (1996). Introductory Mycology (4th ed.). Wiley. ISBN   978-0471522294.
  10. ↑ Dick, M.W. (2013). Straminipilous Fungi: Systematics of the Peronosporomycetes Including Accounts of the Marine Straminipilous Protists, the Plasmodiophorids and Similar Organisms. Springer. ISBN   978-94-015-9733-3.
  11. ↑ "Stramenipila M.W. Dick (2001)". MycoBank. International Mycological Association.
  12. ↑ Patterson, D. J. (1989). "Stramenopiles: chromophytes from a protistological perspective". In Green, J. C.; Leadbeater, B. S. C.; Diver, W. L. (eds.). The chromophyte algae: problems and perspectives. Oxford: Clarendon Press. pp. 357–379.
  13. ↑ Patterson, David J. (1999). "The Diversity of Eukaryotes". The American Naturalist. 154 (S4): S96–S124. doi:10.1086/303287. PMID   10527921. S2CID   4367158.
  14. ↑ David, J. C. (2002). "A preliminary catalogue of the names of fungi above the rank of order". Constancea (83): 1–30.
  15. ↑ Luther, Alexander F. (1899). Über Chlorosaccus eine neue Gattung der Süsswasseralgen nebst einiger Bemerkungen zur Systematik verwandter Algen[About Chlorosaccus a new genus of freshwater algae together with some comments on the systematics of related algae] (in German). Stockholm: Norstedt. pp. 1–22.
  16. ↑ Copeland, H. F. (1956). The Classification of Lower Organisms. Palo Alto, California: Pacific Books.
  17. ↑ Blackwell, W. H. (2009). "Chromista revisited: A dilemma of overlapping putative kingdoms, and the attempted application of the botanical code of nomenclature" (PDF). Phytologia. 91 (2).
  18. ↑ Patterson, Colin (1982). "Morphological characters and homology". In Joysey, Kenneth A.; Friday, A. E. (eds.). Problems in Phylogenetic Reconstruction. Systematics Association Special Volume 21. London: Academic Press. ISBN   978-0-1239-1250-3.
  19. ↑ Hee, Wei Yih; Blackman, Leila M.; Hardham, Adrienne R. (2019). "Characterisation of Stramenopile-specific mastigoneme proteins in Phytophthora parasitica". Protoplasma. 256 (2): 521–535. doi:10.1007/s00709-018-1314-1. PMID   30302550. S2CID   52947780.
  20. ↑ Bouck, G. Benjamin (1 August 1971). "The structure, origin, and composition of the tubular mastigonemes of the Ochromonas flagellum". Journal of Cell Biology. 50 (2): 362–384. doi:10.1083/jcb.50.2.362. PMC   2108286 . PMID   5123323.
  21. ↑ Blackman, Leila M.; Arikawa, Mikihiko; Yamada, Shuhei; Suzaki, Toshinobu; Hardham, Adrienne R. (2011). "Identification of a Mastigoneme Protein from Phytophthora nicotianae". Protist. 162 (1): 100–114. doi:10.1016/j.protis.2010.01.005. PMID   20663714.
  22. ↑ Yoon, H.S.; Andersen, R.A.; Boo, S.M.; Bhattacharya, D. (2009). "Stramenopiles". Encyclopedia of Microbiology. Elsevier. pp. 721–731. doi:10.1016/b978-012373944-5.00253-4. ISBN   9780123739445.
  23. ↑ Fu, Gang; Nagasato, Chikako; Oka, Seiko; Cock, J. Mark; Motomura, Taizo (2014). "Proteomics Analysis of Heterogeneous Flagella in Brown Algae (Stramenopiles)" (PDF). Protist. 165 (5): 662–675. doi:10.1016/j.protis.2014.07.007. PMID   25150613. S2CID   7936118.
  24. ↑ Leipe, D. D.; Wainright, P. O.; Gunderson, J. H.; et al. (1994). "The stramenopiles from a molecular perspective: 16S-like rRNA sequences from Labyrinthuloides minuta and Cafeteria roenbergensis". Phycologia. 33 (5): 369–377. doi:10.2216/i0031-8884-33-5-369.1.
  25. ↑ Leyland, Ben; Leu, Stefan; Boussiba, Sammy (2017). "Are Thraustochytrids algae?". Fungal Biology. 121 (10): 835–840. doi:10.1016/j.funbio.2017.07.006. PMID   28889907.
  26. 1 2 Derelle, Romain; López-García, Purificación; Timpano, Hélène; Moreira, David (10 August 2016). "A Phylogenomic Framework to Study the Diversity and Evolution of Stramenopiles (=Heterokonts)". Molecular Biology and Evolution. 33 (11): 2890–2898. doi:10.1093/molbev/msw168. PMC   5482393 . PMID   27512113.
  27. ↑ Roberts, Tamalee; Stark, Damien; Harkness, John; Ellis, John (2014). "Update on the pathogenic potential and treatment options for Blastocystis sp". Gut Pathogens. 6 (1): 17. doi: 10.1186/1757-4749-6-17 . PMC   4039988 . PMID   24883113.
  28. ↑ Olsen, O. Wilford (1986). Animal Parasites : their life cycles and ecology. New York: Dover. pp. 56, 74–75. ISBN   0-486-65126-6. OCLC   13123309.
  29. ↑ Nowicki, Marcin; et al. (17 August 2011). "Potato and tomato late blight caused by Phytophthora infestans: An overview of pathology and resistance breeding". Plant Disease . 96 (1). American Phytopathological Society: 4–17. doi: 10.1094/PDIS-05-11-0458 . PMID   30731850.
  30. ↑ Yool, Andrew; Tyrrell, Toby (2003). "Role of diatoms in regulating the ocean's silicon cycle". Global Biogeochemical Cycles. 17 (4): n/a. Bibcode:2003GBioC..17.1103Y. CiteSeerX   10.1.1.394.3912 . doi:10.1029/2002GB002018. S2CID   16849373.
  31. ↑ Cock, J. Mark; Peters, Akira F.; Coelho, Susana M. (2011-08-09). "Brown algae". Current Biology. 21 (15): R573–R575. doi: 10.1016/j.cub.2011.05.006 . PMID   21820616.
  32. ↑ Guiry, Wendy (7 April 2011). "Cafeteria T.Fenchel & D.J.Patterson 1988". AlgaeBase. Retrieved 17 March 2023.
  33. ↑ Fenchel, T.; Patterson, D. J. (1988). "Cafeteria roenbergensis nov. gen., nov. sp., a heterotrophic microflagellate from marine plankton". Marine Microbial Food Webs. 3: 9–19.
  34. ↑ Krylov, M. V.; Dobrovolskii, A. A.; Issi, I. V.; Michaelevich, B. I.; Podlipaev, S. A.; Reshetnyak, V. V.; Seravin, L. N.; et al. 1980. New concepts for the system of unicellular organisms. Trudy Zoologischkei Institut Akademiya Nayuk, SSSR 94:122–132.
  35. ↑ Burki, F.; Shalchian-Tabrizi; Pawlowski, J. (August 2008). "Phylogenomics reveals a new 'megagroup' including most photosynthetic eukaryotes". Biology Letters. 4 (4): 366–369. doi:10.1098/rsbl.2008.0224. PMC   2610160 . PMID   18522922.
  36. ↑ Oborník, Miroslav; Lukeš, Julius (2013). "Cell Biology of Chromerids". International Review of Cell and Molecular Biology. Vol. 306. Elsevier. pp. 333–369. doi:10.1016/b978-0-12-407694-5.00008-0. ISBN   9780124076945. PMID   24016529.
  37. ↑ Tikhonenkov, Denis V.; Jamy, Mahwash; Borodina, Anastasia S.; Belyaev, Artem O.; Zagumyonnyi, Dmitry G.; Prokina, Kristina I.; Mylnikov, Alexander P.; Burki, Fabien; Karpov, Sergey A. (2022). "On the origin of TSAR: morphology, diversity and phylogeny of Telonemia". Open Biology. 12 (3). The Royal Society. doi: 10.1098/rsob.210325 . ISSN   2046-2441. PMC   8924772 . PMID   35291881.
  38. 1 2 Cavalier-Smith, Thomas; Scoble, Josephine Margaret (2013). "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.
  39. 1 2 3 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.
  40. 1 2 3 Shiratori, Takashi; Thakur, Rabindra; Ishida, Ken-ichiro (2017). "Pseudophyllomitus vesiculosus (Larsen and Patterson 1990) Lee, 2002, a Poorly Studied Phagotrophic Biflagellate is the First Characterized Member of Stramenopile Environmental Clade MAST-6". Protist. 168 (4): 439–451. doi:10.1016/j.protis.2017.06.004. ISSN   1434-4610. PMID   28822908.
  41. 1 2 Elizabeth J. Weston; Yana Eglit; Alastair G. B. Simpson (6 October 2023). "Kaonashia insperata gen. et sp. nov., a eukaryotrophic flagellate, represents a novel major lineage of heterotrophic stramenopiles". Journal of Eukaryotic Microbiology . doi:10.1111/JEU.13003. ISSN   1066-5234. Wikidata   Q123562228.
  42. ↑ Anna Cho; Denis V. Tikhonenkov; Gordon Lax; Kristina I. Prokina; Patrick J. Keeling (10 November 2023). "Phylogenomic position of genetically diverse phagotrophic stramenopile flagellates in the sediment-associated MAST-6 lineage and a potentially halotolerant placididean". Molecular Phylogenetics and Evolution . 190: 107964. doi:10.1016/J.YMPEV.2023.107964. ISSN   1055-7903. PMID   37951557. Wikidata   Q124719861.
  43. ↑ 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.
  44. ↑ Wetherbee R, Jackson CJ, Repetti SI, Clementson LA, Costa JF, van de Meene A, Crawford S, Verbruggen H (April 2019). "The golden paradox - a new heterokont lineage with chloroplasts surrounded by two membranes". J Phycol. 55 (2): 257–278. doi:10.1111/jpy.12822. hdl: 11343/233613 . PMID   30536815. S2CID   54477112.
  45. ↑ Medlin LK, Desdevises Y (2018). "Sequence analysis confirms a new algal class". Vie et Milieu/Life & Environment.
  46. ↑ Graf, Louis; Yoon, Hwan Su (21 July 2021). "Olisthodiscophyceae, the 17th heterokont algal class". Journal of Phycology. 57 (4): 1091–1093. doi:10.1111/jpy.13184. PMID   34289104. S2CID   236175098.
  47. ↑ Barcytė, Dovilė; Eikrem, Wenche; Engesmo, Anette; Seoane, Sergio; Wohlmann, Jens; Horák, Aleš; Yurchenko, Tatiana; Eliáš, Marek (2 March 2021). "Olisthodiscus represents a new class of Ochrophyta". Journal of Phycology. 57 (4): 1094–1118. doi:10.1111/jpy.13155. hdl: 10852/86515 . PMID   33655496. S2CID   232101186.
  48. ↑ Graf L, Yang EC, Han KY, Küpper FC, Benes KM, Oyadomari JK, Herbert RJH, Verbruggen H, Wetherbee R, Andersen RA, Yoon HS (December 2020). "Multigene Phylogeny, Morphological Observation and Re-examination of the Literature Lead to the Description of the Phaeosacciophyceae Classis Nova and Four New Species of the Heterokontophyta SI Clade". Protist. 171 (6): 125781. doi: 10.1016/j.protis.2020.125781 . PMID   33278705. S2CID   227315556.
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