Overview

Comprehensive Description

Description of Pelomyxa

Free-living polymorphic species from anaerobic or micro-aerophilic habitats, with large multinucleate amoeboid feeding form, up to 5000 microns long, with small non-motile flagella, endosymbiotic bacteria, refringent cytoplasmic granules (sand); movement is directed by fountain-flow cytoplasm and anterior monopodium. Posterior uroid active in food uptake; cysts about 100 microns in diameter also present. Found in freshwater and soils. The cells have many cones of microtubules each arising from the base and sides of a single basal body; sometimes enclosing a nucleus. One to many nuclei, may be surrounded by endosymbiotic bacteria. Cytoplasm contains 2 to several glycogen bodies as large as the nuclei (10 µm). With or without 9+2 arrangement of microtubules in flagellar axoneme. There is no clear synapomorphy, but this genus is distinguished because the most form most usually encountered is a large multinucleate amoeboid stage with many inactive flagella. This currently monotypic genus has been the generic vehicle for 25 nominal species, of which, 18 have been synonymised with Pelomyxa palustris by Whatley and Chapman-Andresen, 5 have been transferred to other genera and 2 are nomina nuda. This species has a widespread distribution. Many of the nominal species assigned to Pelomyxa are regarded different forms (life stages) of one species. A complex life cycle has been described. In spring, cysts release small binucleate amoebae, which grow and become multinucleate and acquire endosymbiotic bacteria. They become elongate with a posterior uroid, and flagella are evident at this stage. The cells later become spherical, and endosymbiotic bacteria congregate around the nuclei. These cells fragment into rosettes giving rise either to cysts in winter or to small amoebae which may undergo another cycle of development. There are endosymbionts that include a methanogenic bacterium, a long bacilliform bacterium resembling Methanobacterium formicium, and a large cylindrical bacterium with a characteristic internal cleft. Type species: Pelomyxa palustris.
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Pelomyxa

Pelomyxa is a genus of giant flagellar amoeboid protists, usually 500-800 μm but occasionally up to 5 mm in length, found in anaerobic or microaerobic bottom sediments of stagnant freshwater ponds or slow-moving streams.[1]

The genus was created by R. Greeff, in 1874, with Pelomyxa palustris as its type species.[2] In the decades following the erection of Pelomyxa, researchers assigned numerous new species to it. However, in the last quarter of the 20th century, investigators reduced the genus to a single species, Pelomyxa palustris, which was understood to be a highly changeable organism with a complex life cycle, whose various phases had been mistaken for separate species.[3][4] All described species were relegated to the status of synonyms, or moved to the unrelated genus Chaos.

Since 2004, four new Pelomyxa species have been described, and two older species have been redescribed and confirmed as valid members of the genus. These developments have raised new questions about the nature of Pelomyxa palustris itself.[1][5][6][7][8][9]

Characteristics[edit]

Pelomyxa have multiple nuclei,[10] which can number from two to several thousand in rare cases. A moving cell is cylindrical in shape, with a single hemispherical pseudopod at the front and a semipermanent projection called a uroid at the back, which is covered in tiny non-motile flagella. They consume a wide variety of food, and have many vacuoles containing both food, such as diatoms, and debris such as sand.

Classification[edit]

The classification of Pelomyxa has been the subject of considerable discussion, in recent decades.

Pelomyxa lack mitochondria, as well as several other organelles usually found in eukaryote cells (notably, peroxisomes and dictyosomes). At one time, they were also believed to lack flagella and to be incapable of mitosis. As nucleated cells that lacked "nearly every other cell-inclusion of eukaryotes",[11] Pelomyxa were, for a time, regarded as surviving "proto-Eukaryotes",[12] standing somewhere between the bacteria and the modern cell. In 1973, it was proposed that the ancestors of Pelomyxa palustris had branched off from the eukaryote line before the advent of mitochondria[13] In 1976, Jean M. Whatley wrote that Pelomyxa palustris "may justly be considered the most primitive eukaryotic organism living today."[14] As such, the organism was potentially a modern analogue of the ancestral eukaryote that, according to the theory of serial endosymbiosis, internalized the bacterial symbiont that later evolved into the mitochondria of the modern cell. The species was known to host several bacterial symbionts. While the function of these was unclear, Whatley argued that they might provide a useful evolutionary example, indicating the "ways in which a bacterial mitochondrial transformation might have been attained."[14]

In 1982, Lynn Margulis created the subclass Caryoblastea (or Pelobiontidae) for "anaerobic ameobas that lack undulipodia," and assigned Pelomyxa to it as the only member of the group.[15] The following year, Cavalier-Smith included the genus with several other "primitive" amitochondriate amoeboids in a new taxonomical group: the Archamoebae.[16] The Archamoebae were, in turn, recruited to the new kingdom of Archezoa, along with other amitochondriate eukaryotes, the Metamonads and the Microsporidia.

The primitivity of Pelomyxa came into doubt in 1988, when Joe I. Griffin published a structural study of Pelomyxa palustris showing that the species does, after all, possess rudimentary flagella, and that it does mitose. Griffin concluded that "Pelomyxa is neither primitive nor different from related forms, once it is realized that its relatives are amoeboid flagellates."[17] In 1995, the case against Pelomyxa's primitivity became stronger still, when molecular analysis revealed that the ancestors of Pelomyxa palustris had most probably possessed mitochondria.[18] By the end of the decade, it was clear that all members of Cavalier-Smith's Archamoebae were descended from mitochondriate cells. In other words, they were not early-branching or "primitive" eukaryotes at all, but rather "degenerate protists" that had lost organelles their ancestors had possessed.[19]

Consequently, Pelomyxa and the other Archamoeba were reassigned to the phylum Amoebozoa, under the subphylum Conosa (shared with the Mycetozoan slime moulds).[20] Kingdom Archezoa was eliminated entirely.[21]

Video gallery[edit]

Sand particles in Pelomyxa cytoplasm

References[edit]

  1. ^ a b Chistyakova, L. V., and A. O. Frolov. "Light and electron microscopic study of Pelomyxa stagnalis sp. n.(Archamoebae, pelobiontida)." Cell and Tissue Biology 5.1 (2011): 90-97.
  2. ^ Greeff, Richard. "Pelomyxa palustris (Pelobius), ein amöberartiger Organismus des süssen Wassers." Archiv für Mikroskopische Anatomie 10.1 (1874): 51-73.
  3. ^ Griffin, Joe I. (1988) "Fine Structure and Taxonomic Position of the Giant Amoeboid Pelomyxa palustris." J. Protozool. 35(2):313
  4. ^ Brugerolle G. and Patterson D. 2000. Order Pelobiontida Page 1976. In: An illustrated guide to the protozoa, second edition (Eds. Lee J., Leedale G. and Bradbury P.). Allen press inc., Lawrence, USA. pp. 1097-1103.
  5. ^ Frolov, Alexander O., Ludmila V. Chystjakova, and Andrew V. Goodkov. "A new pelobiont protist Pelomyxa corona sp. n.(Peloflagellatea, Pelobiontida)." Protistology 3 (2004): 233-241.
  6. ^ Frolov, Alexander O., Ludmila V. Chystjakova, and Andrew V. Goodkov. "Light and electron microscopic study of Pelomyxa binucleata (Gruber, 1884)(Peloflagellatea, Pelo biontida)." Protistology 4 (2005): 57-73.
  7. ^ Frolov, A., et al. "Structure and Development of Pelomyxa gruberi sp. n.(Peloflagellatea, Pelobiontida)." Protistology 4 (2006): 227-244.
  8. ^ Frolov, A. O., et al. "Light and electron microscopic investigation of Pelomyxa prima (Gruber, 1884)(Peloflagellatea, Pelobiontida)." Tsitologiia 47.1 (2005): 89-98.
  9. ^ Frolov, A. O., L. V. Chistiakova, and M. N. Malysheva. "Light-and electron-microscopical study of Pelomyxa flava sp. n.(archamoebae, pelobiontida)]." Tsitologiia 52.9 (2010): 776.
  10. ^ Daniels, E.; Pappas, G. (1994). "Reproduction of nuclei in Pelomyxa palustris". Cell biology international 18 (8): 805–812. doi:10.1006/cbir.1994.1113. PMID 7804157.  edit
  11. ^ Margulis, Lynn (1982, 1988, 1998). Five kingdoms: an illustrated guide to the phyla of life on earth. 4th ed. W. H. Freeman co. p. 120. ISBN 0-7167-3026-X. 
  12. ^ Whatley, Jean M. et al. (April 1979). "From Extracellular to intracellular: the establishment of mitochondria and chloroplasts." Proceedings of the Royal Society of London. Series B, Biological Sciences. 204(155):51)
  13. ^ Bovee, E. C., and T. L. Jahn. 1973. "Taxonomy and phylogeny." Pages 38–76 in K. W. Jeon, ed. The biology of amoeba. Academic Press, New York; cited in Roger, Andrew J. (1999) "Reconstructing Early events in Eukaryotic Evolution." The American Naturalist. 154(S4):S147
  14. ^ a b Whatley, Jean M. (January 1976). "Bacteria and Nuclei in Pelomyxa Palustris: Comments on the Theory of Serial Endosymbiosis". New Phytologist 76 (1): 111–120. doi:10.1111/j.1469-8137.1976.tb01443.x. JSTOR 2431427. 
  15. ^ Margulis, Lynn (1982, 1988, 1998). Five kingdoms: an illustrated guide to the phyla of life on earth. 4th ed. W. H. Freeman co. p. 130. ISBN 0-7167-3026-X. 
  16. ^ Cavalier-Smith, T. 1983. "A 6-kingdom classification and a unified phylogeny." In W. Schwemmler and H. E. A. Schenk (ed.), Endocytobiology II. de Gruyter, Berlin. pp. 1027-1034.
  17. ^ Griffin, Joe I. (1988) "Fine Structure and Taxonomic Position of the Giant Amoeboid Pelomyxa palustris." J. Protozool. 35(2):300-315
  18. ^ Morin, L., and J.-P. Mignot. 1995. Are Archamoebae true Archezoa? the phylogenetic position of Pelomyxa sp. as inferred from large subunit ribosomal RNA sequencing. European Journal of Protistology 31:402.
  19. ^ Edgcomb VP, Simpson AG, Zettler LA, et al. (June 2002). "Pelobionts are degenerate protists: insights from molecules and morphology". Mol. Biol. Evol. 19 (6): 978–82. doi:10.1093/oxfordjournals.molbev.a004157. PMID 12032256. 
  20. ^ T. Cavalier-Smith. A revised six-kingdom system of life. Biological Reviews (1998). 73:203-266.
  21. ^ Keeling, PJ (1998). "A kingdom's progress: Archezoa and the origin of eukaryotes". BioEssays 20 (1): 87–95. doi:10.1002/(sici)1521-1878(199801)20:1<87::aid-bies12>3.0.co;2-4. 
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