Mixotricha paradoxa

Last updated

Mixotricha paradoxa
Mixotricha paradoxa color and.png
Scientific classification
Domain:
(unranked):
Phylum:
Class:
Order:
Family:
Genus:
Mixotricha
Species:
M. paradoxa
Binomial name
Mixotricha paradoxa
Sutherland, 1933

Mixotricha paradoxa is a species of protozoan that lives inside the gut of the Australian termite species Mastotermes darwiniensis .

Contents

It is composed of five different organisms: three bacterial ectosymbionts live on its surface for locomotion and at least one endosymbiont lives inside to help digest cellulose in wood to produce acetate for its host(s).

Mixotricha mitochondria degenerated in hydrogenosomes and mitosomes and lost the ability to produce energy aerobically by oxidative phosphorylation. [1] [2] The mitochondria-derived nuclear genes were however conserved. [2]

Discovery

The name was given by the Australian biologist J.L. Sutherland, who first described Mixotricha in 1933. [3] [4] The name means "the paradoxical being with mixed-up hairs" because this protist has both cilia and flagella, which was not thought to be possible for protists. [5] [2]

Behavior

Mixotricha is a species of protozoan that lives inside the gut of the Australian termite species Mastotermes darwiniensis and has multiple bacterial symbionts. [6] [7]

Mixotricha is a large protozoan .5 millimetres (0.020 in) long and contains hundreds of thousands of bacteria. [5] It is an endosymbiont and digests cellulose for the termite. [5]

Trichomonads like Mixotricha reproduce by a special form of longitudinal fission, leading to large numbers of trophozoites in a relatively short time. Cysts never form, so transmission from one host to another is always based on direct contact between the sites they occupy. [8]

Anatomy

Species of the order Trichomonadida typically have four to six flagella at the cell's apical pole, one of which is recurrent - that is, it runs along a surface wave, giving the aspect of an undulating membrane. Mixotricha paradoxa have four weak flagella that serve as rudders. [9] It has four large flagella at the front end, three pointing forwards and one backward. [5]

The basal bodies are also bacteria, not spirochaetes but oval, pill-shaped bacteria. There is a one-to-one relationship between a bracket, a spirochaete, and a basal bacterium. Each bracket has one spirochaete running through it and one pill bacterium at its base as the basal body. [5] It has not been shown definitely, but the basal bodies could also be making cellulases that digest wood. [5]

Endosymbionts for biochemical processes

At least one endosymbiont lives inside the protist to help digest cellulose and lignin, a major component of the wood the termites eat. The cellulose gets converted to glucose then to acetate, and the lignin is digested directly to acetate. [2] The acetate probably crosses the termite gut membrane to be digested later. [2]

Mixotricha forms a mutualistic relationship with bacteria living inside the termite. There are a total of four species of bacterial symbionts. It has spherical bacteria inside the cell, which function as mitochondria, which Mixotricha lacks. Mixotricha mitochondria degenerated and lost the ability to produce energy aerobically by oxidative phosphorylation. [1] [2] Mitochondrial relics include hydrogenosomes which produce hydrogen and small structures called mitosomes. [2]

Ectosymbionts for movement

Three surface colonising bacteria are anchored on the surface. [10]

The flagella and cilia are actually two different single celled organisms. The ciliate belongs to an archaic group that used to be called archezoa but this term is no longer in fashion. [11] It has four weak flagella, which serve as a rudder. [12] [11]

While Mixotricha has four anterior flagella, it does not use them for locomotion, but more for steering. [5] For locomotion, about 250,000 hairlike Treponema spirochaetes, a species of helical bacteria, are attached to the cell surface and provide the cell with cilia-like movements. [2]

The wavelength of the cilia is about .1 millimetres (0.0039 in) and suggests that the spirochaetes are somehow in touch with each other. [5]

Mixotricha also has rod-shaped bacteria arranged in an ordered pattern on the surface of the cell. [13]

Each spirochaete has its own little emplacement, called a 'bracket'. [14] Spirochetes move continuously forwards or backwards but when they are attached they move in one direction. [2]

Sperm tails might have their origin in spirochaetes. [2] The evidence that cilia (undulipodia) are symbiotic bacteria is found unpersuasive. [5]

Genome

Mixotricha have five genomes, as they form very close symbiotic relationships with four types of bacteria. [15] It is a good example organism for symbiogenesis and nestedness. [2]

There are two spirochete and one-rod bacteria on its surface, one endosymbiotic bacteria inside to digest cellulose and the host nucleus. [2]

Related Research Articles

<span class="mw-page-title-main">Endosymbiont</span> Organism that lives within the body or cells of another organism

An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualistic relationship. Examples are nitrogen-fixing bacteria, which live in the root nodules of legumes, single-cell algae inside reef-building corals, and bacterial endosymbionts that provide essential nutrients to insects.

The evolution of flagella is of great interest to biologists because the three known varieties of flagella – each represent a sophisticated cellular structure that requires the interaction of many different systems.

<span class="mw-page-title-main">Unicellular organism</span> Organism that consists of only one cell

A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and eukaryotic organisms. Most prokaryotes are unicellular and are classified into bacteria and archaea. Many eukaryotes are multicellular, but some are unicellular such as protozoa, unicellular algae, and unicellular fungi. Unicellular organisms are thought to be the oldest form of life, with early protocells possibly emerging 3.5–4.1 billion years ago.

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

The parabasalids are a group of flagellated protists within the supergroup Excavata. Most of these eukaryotic organisms form a symbiotic relationship in animals. These include a variety of forms found in the intestines of termites and cockroaches, many of which have symbiotic bacteria that help them digest cellulose in woody plants. Other species within this supergroup are known parasites, and include human pathogens.

<span class="mw-page-title-main">Metamonad</span> Phylum of excavate protists

The metamonads are a large group of flagellate amitochondriate microscopic eukaryotes. They include the retortamonads, diplomonads, parabasalids, oxymonads, and a range of more poorly studied taxa, most of which are free-living flagellates. All metamonads are anaerobic, and most members of the four groups listed above are symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals.

<i>Reticulitermes flavipes</i> Species of insect found in North America

Reticulitermes flavipes, the eastern subterranean termite, is the most common termite found in North America. These termites are the most economically important wood destroying insects in the United States and are classified as pests. They feed on cellulose material such as the structural wood in buildings, wooden fixtures, paper, books, and cotton. A mature colony can range from 20,000 workers to as high as 5 million workers and the primary queen of the colony lays 5,000 to 10,000 eggs per year to add to this total.

<span class="mw-page-title-main">Trichomonadida</span> Order of flagellated protists

Trichomonadida is an order of anaerobic protists, included with the parabasalids. Members of this order are referred to as trichomonads.

Symbiotic bacteria are bacteria living in symbiosis with another organism or each other. For example, rhizobia living in root nodules of legumes provide nitrogen fixing activity for these plants.

<i>Trichonympha</i> Genus of flagellated protists

Trichonympha is a genus of single-celled, anaerobic parabasalids of the order Hypermastigia that is found exclusively in the hindgut of lower termites and wood roaches. Trichonympha’s bell shape and thousands of flagella make it an easily recognizable cell. The symbiosis between lower termites/wood roaches and Trichonympha is highly beneficial to both parties: Trichonympha helps its host digest cellulose and in return receives a constant supply of food and shelter. Trichonympha also has a variety of bacterial symbionts that are involved in sugar metabolism and nitrogen fixation.

<i>Mastotermes darwiniensis</i> Species of termite

Mastotermes darwiniensis, common names giant northern termite and Darwin termite, is a termite species found only in northern Australia. It is the most primitive extant termite species. Contrary to common belief, this species does not form mounds as the nests are subterranean and inconspicuous. Colonies will readily occupy and infest decomposing wood but primarily live in a complex subterranean network of tunnels and galleries which they use to travel to new food sites. Colonies may eventually split and form isolated satellite colonies.

<span class="mw-page-title-main">Protozoan infection</span> Parasitic disease caused by a protozoan

Protozoan infections are parasitic diseases caused by organisms formerly classified in the kingdom Protozoa. These organisms are now classified in the supergroups Excavata, Amoebozoa, Harosa, and Archaeplastida. They are usually contracted by either an insect vector or by contact with an infected substance or surface.

<span class="mw-page-title-main">Malawimonadidae</span> Family of protists

Malawimonadidae is a family of unicellular eukaryotes of outsize importance in understanding eukaryote phylogeny.

Trimastix is a genus of excavate protists, the sole occupant of the order Trimastigida. Trimastix are bacterivorous, free living and anaerobic. It was first observed in 1881 by William Kent. There are few known species, and the genus's role in the ecosystem is largely unknown. However, it is known that they generally live in marine environments within the tissues of decaying organisms to maintain an anoxic environment. Much interest in this group is related to its close association with other members of Preaxostyla. These organisms do not have classical mitochondria, and as such, much of the research involving these microbes is aimed at investigating the evolution of mitochondria.

Psalteriomonas is a genus of excavates in the group of Heterolobosea. The genus was first discovered and named in 1990. It contains amoeboflagellate cells that live in freshwater anaerobic sediments all over the world. The microtubule-organizing ribbon and the associated microfibrillar bundles of the mastigote system is the predominant feature in Psalteriomonas. This harp-shaped complex gives rise to the name of this genus. Psalteriomonasforms an endosymbiotic relationship with methanogenic bacteria, especially with Methanobacterium formicicum There are currently three species in this genus: P. lanterna, P. vulgaris, and P. magna.

Oxymonas is a genus of Excavata.

<i>Mastotermes</i> Genus of termites

Mastotermes is a genus of termites. The sole living species is Mastotermes darwiniensis, found only in northern Australia. A number of extinct taxa are known from fossils. It is a very peculiar insect, the most primitive termite alive. As such, it shows notable similarities to cockroaches in the family Cryptocercidae, the termites' closest relatives. These similarities include the anal lobe of the wing and the laying of eggs in bunches, rather than singly. The termites were traditionally placed in the Exopterygota, but such an indiscriminate treatment makes that group a paraphyletic grade of basal neopterans. Thus, the cockroaches, termites and their relatives are nowadays placed in a clade called Dictyoptera.

<i>Angomonas deanei</i> Species of parasitic flagellate protist in the Kinetoplastea class

Angomonas deanei is a flagellated trypanosomatid protozoan. As an obligate parasite, it infects the gastrointestinal tract of insects, and is in turn a host to symbiotic bacteria. The bacterial endosymbiont Ca. "Kinetoplastibacterium crithidii" maintains a permanent mutualistic relationship with the protozoan such that it is no longer able to reproduce and survive on its own. The symbiosis, subsequently also discovered in varying degrees in other protists such as Strigomonas culicis, Novymonas esmeraldas, Diplonema japonicumand Diplonema aggregatum are considered as good models for the understanding of the evolution of eukaryotes from prokaryotes, and on the origin of cell organelles.

Monocercomonoides is a genus of flagellate Excavata belonging to the order Oxymonadida. It was established by Bernard V. Travis and was first described as those with "polymastiginid flagellates having three anterior flagella and a trailing one originating at a single basal granule located in front of the anteriorly positioned nucleus, and a more or less well-defined axostyle". It is the first eukaryotic genus to be found to completely lack mitochondria, and all hallmark proteins responsible for mitochondrial function. The genus also lacks any other mitochondria related organelles (MROs) such as hydrogenosomes or mitosomes. Data suggests that the absence of mitochondria is not an ancestral feature, but rather due to secondary loss. Monocercomonoides sp. was found to obtain energy through an enzymatic action of nutrients absorbed from the environment. The genus has replaced the iron-sulfur cluster assembly pathway with a cytosolic sulfur mobilization system, likely acquired by horizontal gene transfer from a eubacterium of a common ancestor of oxymonads. These organisms are significant because they undermine assumptions that eukaryotes must have mitochondria to properly function. The genome of Monocercomonoides exilis has approximately 82 million base pairs, with 18 152 predicted protein-coding genes.

Lemuel Roscoe Cleveland was an American zoologist and protistologist, famous for giving the first, strong empirical proof for the existence of a symbiotic relationship between internal microorganisms and their metazoan host.

<span class="mw-page-title-main">Marine protists</span> Protists that live in saltwater or brackish water

Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as marine single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics because they are paraphyletic.

References

  1. 1 2 Cepicka, Ivan; Dolan, Michael F.; Gile, Gillian (1 September 2017). "Parabasalia". Handbook of the Protists. pp. 1175–1218. doi:10.1007/978-3-319-28149-0_9. ISBN   978-3-319-28147-6.{{cite book}}: |journal= ignored (help)
  2. 1 2 3 4 5 6 7 8 9 10 11 12 Thompson, William Irwin (1991). Gaia 2: Emergence : The New Science of Becoming. SteinerBooks. pp. 51–58. ISBN   9780940262409.
  3. Jean L. Sutherland: Protozoa from Australian Termites. Quarterly Journal of Microscopical Science, Band s2-76, S. 145-173. (Abstract)
  4. L. R. Cleveland, A. V. Grimstone: The Fine Structure of the Flagellate Mixotricha paradoxa and Its Associated Micro-Organisms. Proceedings of the Royal Society of London. Series B, Biological Sciences, Band 159, 1964, S. 668-686. doi : 10.1098/rspb.1964.0025
  5. 1 2 3 4 5 6 7 8 9 Dawkins, Richard; Wong, Yan (2016). The Ancestor's Tale. Houghton Mifflin Harcourt. ISBN   978-0544859937.
  6. Radek R, Nitsch G (November 2007). "Ectobiotic spirochetes of flagellates from the termite Mastotermes darwiniensis: attachment and cyst formation". Eur. J. Protistol. 43 (4): 281–94. doi:10.1016/j.ejop.2007.06.004. PMID   17764914.
  7. Brugerolle G (October 2004). "Devescovinid features, a remarkable surface cytoskeleton, and epibiotic bacteria revisited in Mixotricha paradoxa, a parabasalid flagellate". Protoplasma. 224 (1–2): 49–59. doi:10.1007/s00709-004-0052-8. PMID   15726809. S2CID   21431174.
  8. Kamaruddin, Mudyawati; Tokoro, Masaharu; Rahman, Md. Moshiur; Arayama, Shunsuke; Hidayati, Anggi P.N.; Syafruddin, Din; Asih, Puji B.S.; Yoshikawa, Hisao; Kawahara, Ei (2014). "Molecular Characterization of Various Trichomonad Species Isolated from Humans and Related Mammals in Indonesia". The Korean Journal of Parasitology. 52 (5): 471–478. doi:10.3347/kjp.2014.52.5.471. PMC   4210728 . PMID   25352694.
  9. Overmann, Jörg (10 January 2006). Molecular Basis of Symbiosis . Springer. pp.  76–95. ISBN   9783540282105.
  10. "Movement symbiosis2". www.microbiological-garden.net. Retrieved 27 May 2019.
  11. 1 2 König, Helmut (2006). Intestinal Microorganisms of Termites and Other Invertebrates . Springer. pp.  261–263. ISBN   9783540281801.
  12. "Movement symbiosis". www.microbiological-garden.net. Retrieved 27 May 2019.
  13. Rosenberg, Eugene; Gophna, Uri (30 August 2011). Beneficial Microorganisms in Multicellular Life Forms . Springer. p.  9. ISBN   9783642216800.
  14. Hongoh, Y.; Sato, T.; Dolan, M. F.; Noda, S.; Ui, S.; Kudo, T.; Ohkuma, M. (2007). "The Motility Symbiont of the Termite Gut Flagellate Caduceia versatilis Is a Member of the "Synergistes" Group". Applied and Environmental Microbiology. 73 (19): 6270–6276. Bibcode:2007ApEnM..73.6270H. doi:10.1128/AEM.00750-07. PMC   2074993 . PMID   17675420.
  15. Margulis, Lynn; Sagan, Dorion (June 2001). "The Beast with Five Genomes". Natural History. Archived from the original on 15 November 2006. Retrieved 3 May 2007.