Euglenid

Euglenids
Temporal range: Eocene (53.5Ma) - recent[1]
Euglena viridis, by Ehrenberg
Scientific classification
Domain: Eukaryota
(unranked): Excavata
Phylum: Euglenophyta
Pascher, 1931
Class: Euglenophyceae
Schoenichen, 1925
Major groups[2]

Phototrophs (in general)
   Euglenales/Euglenina
   Eutreptiales/Eutreptiina
   Euglenamorphales/Euglenamorphina
Osmotrophs
   Rhabdomonadales/Rhabdomonadina
Phagotrophs
   ?Heteronematales/Heteronematina
   ?Sphenomonadales/Sphenomonadina

Synonyms
  • Euglenoidina Bütschli, 1884, Blochmann, 1886
  • Euglenoidea Lankester, 1885
  • Euglenida Stein, 1878 sensu Simpson, 1997
  • Euglenoida Cavalier-Smith, 1993

Euglenids (euglenoids, or euglenophytes, formally Euglenida/Euglenoida, ICZN, or Euglenophyceae, ICBN) are one of the best-known groups of flagellates, which are excavate eukaryotes of the phylum Euglenophyta and their cell structure is typical of that group. They are commonly found in freshwater, especially when it is rich in organic materials, with a few marine and endosymbiotic members. Most euglenids are unicellular. Many euglenids have chloroplasts and produce their own food through photosynthesis, but others feed by phagocytosis, or strictly by diffusion. This group is known to contain the carbohydrate paramylon.

Euglenids are said to descend from an ancestor that took up green algae by secondary endosymbiosis.[3]

Structure

Euglenids are distinguished mainly by the presence of a pellicle (periplast). Within its taxon, the pellicle is one of the euglenids' most diverse features from a morphological standpoint.[4] The pellicle is composed of proteinaceous strips underneath the cell membrane, supported by dorsal and ventral microtubules. This varies from rigid to flexible, and gives the cell its shape, often giving it distinctive striations. In many euglenids the strips can slide past one another, causing an inching motion called metaboly. Otherwise they move using their flagella.

Classification

1—2. Ascoglena sp. (Euglenales);
3—4. Cryptoglena sp. (idem);
5—9, 14—15, 24—25, 27-29. Trachelomonas spp. (id.);
10. Eutreptia sp. (Eutreptiales);
11, 20. Astasia spp. (Euglenales);
12. Distigma sp. (Eutreptiales);
13. Menoid[i]um sp. (Rhabdomonadales);
16—18. Colacium sp. (Euglenales);
19, 26. Petalomonas spp. (Sphenomonadales);
21. Sphenomonas sp. (id.);
22—23. Euglenopsis sp. (Euglenales);
30. Peranema sp. (Heteronematales)

The euglenids were first defined by Otto Bütschli in 1884 as the flagellate order Euglenida, as an animal. Botanists subsequently created the algal division Euglenophyta; thus they were classified as both animals and plants, as they share characteristics with both. Conflicts of this nature are exemplary of why the kingdom Protista was adopted. However, they retained their double-placement until the flagellates were split up, and both names are still used to refer to the group. Their chlorophyll are not masked with accessory pigments.

Nutrition

The classification of euglenids is still variable, as groups are being revised to conform with their molecular phylogeny. Classifications have fallen in line with the traditional groups based on differences in nutrition and number of flagella; these provide a starting point for considering euglenid diversity. Different characteristics of the euglenids' pellicles can provide insight into their modes of movement and nutrition.[5]

As with other Euglenozoa, the primitive mode of nutrition is phagocytosis. Prey such as bacteria and smaller flagellates are ingested through a cytostome, supported by microtubules. These are often packed together to form two or more rods, which function in ingestion, and in Entosiphon form an extendable siphon. Most phagotrophic euglenids have two flagella, one leading and one trailing. The latter is used for gliding along the substrate. In some, such as Peranema, the leading flagellum is rigid and beats only at its tip.

Osmotrophic euglenoids

Osmotrophic euglenids are euglenids which have undergone osmotrophy.

Due to a lack of characteristics which are useful for taxonomical purposes, the origin of osmotrophic euglenids is unclear. Though, certain morphological characteristics reveal a small fraction of osmotrophic euglenids are derived from phototrophic and phagotrophic ancestors.[6]

A prolonged absence of light or exposure to harmful chemicals may cause atrophy and absorption of the chloroplasts without otherwise harming the organism. There are a number of species where a chloroplast's absence was formerly marked with separate genera such as Astasia (colourless Euglena) and Hyalophacus (colourless Phacus). Due to the lack of a developed cytostome, these forms feed exclusively by osmotrophic absorption.

Reproduction

Although euglenids share several common characteristics with animals, which is why they were originally classified as so, there is no evidence of euglenids ever using sexual reproduction. This is one of the reasons they could no longer be classified as animals.

For euglenids to reproduce, asexual reproduction takes place in the form of binary fission, and the cells replicate and divide during mitosis and cytokinesis. This process occurs in a very distinct order. First, the basal bodies and flagella replicate, then the cytostome and microtubules (the feeding apparatus), and finally the nucleus and remaining cytoskeleton. Once this occurs, the organism begins to cleave at the basal bodies, and this cleavage line moves towards the center of the organism until two separate euglenids are evident.[7] Because of the way that this reproduction takes place and the axis of separation, this is called longitudinal cell division or longitudinal binary fission.[8]

References

  1. Lee, R.E. (2008). Phycology, 4th edition. Cambridge University Press. ISBN 978-0-521-63883-8.
  2. Leedale, G. F. (1967), Euglenoid Flagellates. Prentice Hall, Englewood Cliffs, 242 p., .
  3. Keeling PJ (2009). "Chromalveolates and the evolution of plastids by secondary endosymbiosis". J. Eukaryot. Microbiol. 56 (1): 1–8. doi:10.1111/j.1550-7408.2008.00371.x. PMID 19335769.
  4. Leander, Brian S.; Farmer, Mark A. (2001-03-01). "Comparative Morphology of the Euglenid Pellicle. II. Diversity of Strip Substructure". Journal of Eukaryotic Microbiology. 48 (2): 202–217. doi:10.1111/j.1550-7408.2001.tb00304.x. ISSN 1550-7408.
  5. Leander, Brian Scott (May 2001). "Evolutionary morphology of the euglenid pellicle". University Of Georgia Theses and Dissertations.
  6. Busse, Ingo; Preisfeld, Angelika (14 April 2018). "Systematics of primary osmotrophic euglenids: a molecular approach to the phylogeny of Distigma and Astasia (Euglenozoa)". International Journal of Systematic and Evolutionary Microbiology. 53 (2): 617–624. doi:10.1099/ijs.0.02295-0. Retrieved 14 April 2018 via Microbiology Society Journals.
  7. "Euglenida". tolweb.org. Retrieved 2017-03-30.
  8. "Reproduction". Euglena. Retrieved 2017-03-31.

Bibliography

  • Ciugulea, I. & Triemer, R. E. (2010) A Color Atlas of Photosynthetic Euglenoids. Michigan State University Press, East Lansing, MI, 204 p., .
  • Leander, B. S., Triemer, R. E., & Farmer, M. A. (2001). Character evolution in heterotrophic euglenids. European Journal of Protistology, 37(3), 337-356, .
  • Leander, B.S., Lax, G., Karnkowska, A., Simpson, A.G.B. (2017). Euglenida. In: Archibald, J.M., Simpson, A.G.B., Slamovits, C. (Eds.). Handbook of the Protists. Springer, pp. 1–42. doi:10.1007/978-3-319-32669-6_13-1
  • Leedale, G. F. (1978). Phylogenetic criteria in euglenoid flagellates. BioSystems 10: 183–187, .
  • Wołowski, K & Hindák, F. (2005). Atlas of Euglenophytes. Cracow: VEDA Publishing House of the Slovak Academy of Sciences, 136 p., .
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