Euglenid

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Euglenids
Temporal range: Eocene (53.5Ma) - recent [1]
Ehrenberg euglena viridis.jpg
Euglena viridis , by Ehrenberg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Discoba
Superphylum: Discicristata
Phylum: Euglenozoa
Class: Euglenophyceae
Schoenichen, 1925
Major groups [2]

Phototrophs (in general)
    Euglenales/Euglenina
    Eutreptiales/Eutreptiina
    Euglenamorphales/Euglenamorphina
Chemotrophs

Contents

    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. Many euglenids feed by phagocytosis, or strictly by diffusion. A monophyletic group consisting of the mixotrophic Rapaza viridis (1 species) and the two groups Eutreptiales (24 species) and Euglenales (983 species) have chloroplasts and produce their own food through photosynthesis. [3] [4] [5] This group is known to contain the carbohydrate paramylon.

Euglenids split from other Euglenozoa more than a billion years ago. The plastids in all extant photosynthetic species is the result from secondary endosymbiosis between a phagotrophic eukaryovorous euglenid and a Pyramimonas -related green alga. [6] In January 2024, some fossilized forms of euglenid had been found to be mistakenly classified as Pseudoschizaea shells. [7]

Structure

Euglenid Body Plan (by Patrick Keeling and Yana Eglit) Euglenid body plan.jpg
Euglenid Body Plan (by Patrick Keeling and Yana Eglit)

Euglenoids are distinguished mainly by the presence of a type of cell covering called a pellicle. Within its taxon, the pellicle is one of the euglenoids' most diverse morphological features. [8] 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

Diagnostic drawings of euglenids Euglenophyceae004.jpg
Diagnostic drawings of euglenids
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) Algen I. (Schizophyceen, Flagellaten, Peridineen) (1910) (17950163265).jpg
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 chlorophylls 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. [9]

As with other Euglenozoa, the primitive mode of nutrition is phagocytosis. Prey such as bacteria and smaller flagellates is 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 that 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. [10]

A prolonged absence of light or exposure to harmful chemicals may cause atrophy and absorption of the chloroplasts without otherwise harming the organism. A number of species exists 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, no evidence has been found of euglenids ever using sexual reproduction. This is one of the reasons they could no longer be classified as animals.[ dubious ]

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. [11] Because of the way that this reproduction takes place and the axis of separation, it is called longitudinal cell division or longitudinal binary fission. [12]

Related Research Articles

<span class="mw-page-title-main">Euglenozoa</span> Phylum of protozoans

Euglenozoa are a large group of flagellate Discoba. They include a variety of common free-living species, as well as a few important parasites, some of which infect humans. Euglenozoa are represented by four major groups, i.e., Kinetoplastea, Diplonemea, Euglenida, and Symbiontida. Euglenozoa are unicellular, mostly around 15–40 μm (0.00059–0.00157 in) in size, although some euglenids get up to 500 μm (0.020 in) long.

<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, the term "flagellate" is included in other terms which are more formally characterized.

<i>Euglena</i> Genus of unicellular flagellate eukaryotes

Euglena is a genus of single cell flagellate eukaryotes. It is the best known and most widely studied member of the class Euglenoidea, a diverse group containing some 54 genera and at least 200 species. Species of Euglena are found in fresh water and salt water. They are often abundant in quiet inland waters where they may bloom in numbers sufficient to color the surface of ponds and ditches green (E. viridis) or red (E. sanguinea).

<span class="mw-page-title-main">Paramylon</span> Chemical compound

Paramylon is a carbohydrate similar to starch. The chloroplasts found in Euglena contain chlorophyll which aids in the synthesis of carbohydrates to be stored as starch granules and paramylon. Paramylon is made in the pyrenoids of Euglena. The euglenoids have chlorophylls a and b and they store their photosynthate in an unusual form called paramylon starch, a β-1,3 polymer of glucose. The paramylon is stored in rod like bodies throughout the cytoplasm, called paramylon bodies, which are often visible as colorless or white particles in light microscopy. Their shape is often characteristic of the Euglena species that produces them.

<i>Euglena gracilis</i> Species of single-celled Eukaryote algae

Euglena gracilis is a freshwater species of single-celled alga in the genus Euglena. It has secondary chloroplasts, and is a mixotroph able to feed by photosynthesis or phagocytosis. It has a highly flexible cell surface, allowing it to change shape from a thin cell up to 100 µm long to a sphere of approximately 20 µm. Each cell has two flagella, only one of which emerges from the flagellar pocket (reservoir) in the anterior of the cell, and can move by swimming, or by so-called "euglenoid" movement across surfaces. E. gracilis has been used extensively in the laboratory as a model organism, particularly for studying cell biology and biochemistry.

<i>Trachelomonas</i> Genus of euglenoids

Trachelomonas is a genus of swimming, free-living euglenoids characterized by the presence of a shell-like covering called a lorica. Details of lorica structure determine the classification of distinct species in the genus. The lorica can exist in spherical, elliptical, cylindrical, and pyriform (pear-shaped) forms. The lorica surface can be smooth, punctuate or striate and range from hyaline, to yellow, or brown. These colors are due to the accumulation of ferric hydroxide and manganic oxide deposited with the mucilage and minerals that comprise the lorica. In Trachelomonas, the presence of a lorica obscures cytoplasmic details of the underlying cell. In each Trachelomonas cell, there is a gap at the apex of the lorica from which the flagellum protrudes. Thickening around this gap results in a rim-like or collar-like appearance. During asexual reproduction, the nucleus divides yielding two daughter cells one of which exits through the opening in the lorica. This new cell then synthesizes its own new lorica.

<span class="mw-page-title-main">Euglenales</span> Order of flagellate eukaryotes

Euglenales is an order of flagellates in the phylum Euglenozoa. The family includes the most well-known euglenoid genus, Euglena, as well as other common genera like Phacus and Lepocinclis.

<span class="mw-page-title-main">Euglenaceae</span> Family of flagellate eukaryotes

Euglenaceae is a family of flagellates in the phylum Euglenozoa. The family includes the most well-known euglenoid genus, Euglena.

<span class="mw-page-title-main">Protozoa</span> Single-celled eukaryotic organisms that feed on organic matter

Protozoa are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically, protozoans were regarded as "one-celled animals".

<i>Colpodella</i> Genus of single-celled organisms

Colpodella is a genus of alveolates comprising 5 species, and two further possible species: They share all the synapomorphies of apicomplexans, but are free-living, rather than parasitic. Many members of this genus were previously assigned to a different genus - Spiromonas.

<i>Phacus</i> Genus of algae

Phacus is a genus of unicellular excavates, of the phylum Euglenozoa, characterized by its flat, leaf-shaped structure, and rigid cytoskeleton known as a pellicle. These eukaryotes are mostly green in colour, and have a single flagellum that extends the length of their body. They are morphologically very flat, rigid, leaf-shaped, and contain many small discoid chloroplasts.

<i>Peranema</i> Genus of protozoans

Peranema is a genus of free-living phagotrophic euglenids. There are more than 20 nominal species, varying in size between 8 and 200 micrometers. Peranema cells are gliding flagellates found in freshwater lakes, ponds and ditches, and are often abundant at the bottom of stagnant pools rich in decaying organic material. Although they belong to the class Euglenoidea, and are morphologically similar to the green Euglena, Peranema have no chloroplasts, and do not conduct autotrophy. Instead, they capture live prey, such as yeast, bacteria and other flagellates, consuming them with the help of a rigid feeding apparatus called a "rod-organ." Unlike the green euglenids, they lack both an eyespot (stigma), and the paraflagellar body (photoreceptor) that is normally coupled with that organelle. However, while Peranema lack a localized photoreceptor, they do possess the light-sensitive protein rhodopsin, and respond to changes in light with a characteristic "curling behaviour."

<i>Euglena sanguinea</i> Species of single cell flagellate eukaryotes

Euglena sanguinea is a species of the genus Euglena. The red colour is due to the presence of astaxanthin and the cells can be populous enough to colour water red. The pigment is used to protect the chloroplasts from light that is too intense, but as the light levels change the cells can take on a green colour as the red pigment is moved to the centre of the cells. Euglena sanguinea is known to make the potent icthyotoxin euglenophycin. Icthyotoxin euglenophycin is a toxin that is very similar in structure to solensopsin, a alkaloid that is found in fire ant venom. This is the only known species of euglenids that is able to form toxic blooms that cause tangible losses to fish farms.

<span class="mw-page-title-main">Diplonemidae</span> Family of protozoans

Diplonemidae is a family of biflagellated unicellular protists that may be among the more diverse and common groups of planktonic organisms in the ocean. Although this family is currently made up of three named genera; Diplonema, Rhynchopus, and Hemistasia, there likely exist thousands of still unnamed genera. Organisms are generally colourless and oblong in shape, with two flagella emerging from a subapical pocket. They possess a large mitochondrial genome composed of fragmented linear DNA. These non-coding sequences must be massively trans-spliced, making it one of the most complicated post-transcriptional editing process known to eukaryotes.

Profª. Dra. Visitación Teresa Dora Conforti de Marconi is an Argentine biologist, algologist, botanist, taxonomist and ecologist. She is a professor in the Department of Biodiversity and Experimental Biology in the University of Buenos Aires in Buenos Aires. She is noted for her numerous taxonomic studies of Euglenophyta, including in polluted rivers in Argentina. Camaleão Lake in Brazil in 1994 and the Caura River in Venezuela in the late 1990s.

Petalomonas is a genus of phagotrophic, flagellated euglenoids. Phagotrophic euglenoids are one of the most important forms of flagellates in benthic aquatic systems, playing an important role in microbial food webs. The traits that distinguish this particular genus are highly variable, especially at higher taxa. However, general characteristics such as a rigid cell shape and single emergent flagellum can describe the species among this genus.

Heteronema is a genus of phagotrophic, flagellated euglenoids that are most widely distributed in fresh water environments. This genus consists of two very distinguishable morphogroups that are phylogenetically closely related. These morphogroups are deciphered based on shape, locomotion and other ultrastructural traits. However, this genus does impose taxonomic problems due to the varying historical descriptions of Heteronema species and its similarity to the genus Paranema. The species H. exaratum, was the first heteronemid with a skidding motion to be sequenced, which led to the discovery that it was not closely related to H. scaphrum, contrary to what was previously assumed, but instead to a sister group of primary osmotrophs. This suggests that skidding heteronemids can also be distinguished phylogenetically, being more closely related to Anisoma, Dinema and Aphageae, than to other species within Heteronema.

Cryptoglena(/ˌkɹɪptoʊˈgliːnə/) is a genus of photosynthetic euglenids that was first described in 1831 by Christian Gottfried Ehrenberg. Today, its circumscription is controversial: Bicudo and Menezes consider twenty-one species as Cryptoglena, of which, nine are uncertain. Cryptoglena species are water-based, living in both freshwater and marine environments. They are biflagellated, with one internal flagellum and one external flagellum, which allows movement through environments as demonstrated by Kim and Shin in the species C. pigra. The cells of Cryptoglena resemble a coffee bean, as they have a groove that runs the length of the cell on one side and makes them U-shaped in cross section. They are ovoid in shape and are small, with the larger cells being on average 25 x 15 μm. After being first described in 1831, little work was done on the genus until the late 1970s and early 1980s, after the scanning electron microscope completed development and was implemented into laboratories. Work then proceeded with the developments of molecular biology, which allows for classifications based on DNA sequences. For Cryptoglena the main DNA used for classification are small subunit (SSU) and large subunit (LSU) rDNA.

Postgaardia is a proposed basal clade of flagellate Euglenozoa, following Thomas Cavalier-Smith. As of April 2023, the Interim Register of Marine and Nonmarine Genera treats the group as a subphylum. A 2021 review of Euglenozoa places Cavalier-Smith's proposed members of Postgaardia in the class Symbiontida. As Euglenozoans may be basal eukaryotes, the Postgaardia may be key to studying the evolution of Eukaryotes, including the incorporation of eukaryotic traits such as the incorporation of alphaproteobacterial mitochondrial endosymbionts.

<i>Urceolus</i> Genus of flagellates

Urceolus is a genus of heterotrophic flagellates belonging to the Euglenozoa, a phylum of single-celled eukaryotes or protists. Described by Russian biologist Konstantin Mereschkowsky in 1877, its type species is Urceolus alenizini. Species of this genus are characterized by deformable sack-shaped cells that exhibit at least one flagellum that is active at the tip. They are found in a variety of water body sediments across the globe. Molecular phylogenies show they belong to a group known as peranemids, closely related to the euglenophyte algae.

References

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  2. Leedale, G. F. (1967). Euglenoid Flagellates. Prentice Hall, Englewood Cliffs.
  3. Karnkowska, Anna; Bennett, Matthew S.; Triemer, Richard E. (2018). "Dynamic evolution of inverted repeats in Euglenophyta plastid genomes". Scientific Reports. 8: 16071. Bibcode:2018NatSR...816071K. doi:10.1038/s41598-018-34457-w. PMID   30375469.
  4. Yoshihisa Hirakawa (2017). Secondary Endosymbioses. Academic Press. p. 323. ISBN   9780128026809.
  5. "Algaebase :: Subclass: Euglenophycidae". Archived from the original on 2020-07-13. Retrieved 2019-10-27.
  6. Zakryś, B; Milanowski, R; Karnkowska, A (2017). "Evolutionary Origin of Euglena". Euglena: Biochemistry, Cell and Molecular Biology. Advances in Experimental Medicine and Biology. Vol. 979. pp. 3–17. doi:10.1007/978-3-319-54910-1_1. ISBN   978-3-319-54908-8. PMID   28429314.
  7. Cassella, Carly (21 January 2024). "Bizarre Fossils Are Neither Plant Nor Animal, But a 'Weird Fusion' of Life". ScienceAlert . Archived from the original on 22 January 2024. Retrieved 22 January 2024.
  8. 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. PMID   12095109. S2CID   2109559.
  9. Leander, Brian Scott (May 2001). "Evolutionary morphology of the euglenid pellicle". University of Georgia Theses and Dissertations.
  10. 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 . PMID   12710635.
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  12. "Reproduction". Euglena. Retrieved 2017-03-31.

Bibliography