Syssomonas | |
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SEM image of Syssomonas. ac = acroneme, fl = flagellum | |
Scientific classification | |
Domain: | Eukaryota |
(unranked): | Holozoa |
Clade: | Pluriformea |
Class: | Corallochytrea |
Order: | Corallochytrida |
Family: | Syssomonadidae Cavalier-Smith 2021 |
Genus: | Syssomonas Tikhonenkov et al. 2017 |
Type species | |
Syssomonas multiformis [1] Tikhonenkov et al. 2017 | |
Type strain | |
Colp-12 MI-PR205 [a] | |
Species | |
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Syssomonas is a monotypic genus of unicellular flagellated protists containing the species Syssomonas multiformis. It is a member of Pluriformea inside the lineage of Holozoa, a clade containing animals and their closest protistan relatives. It lives in freshwater habitats. It has a complex life cycle that includes unicellular amoeboid and flagellated phases, as well as multicellular aggregates, depending on the growth medium and nutritional state.
Syssomonas multiformis is a species of unicellular protists with naked cells (lacking any shell or scales) that presents with a variety of life forms during their complex life cycle. These forms include: round flagellate cells (7–14 μm in diameter) with one posterior flagellum, amoeboflagellate (i.e. with both flagella and pseudopodia) cells, amoeboid non-flagellar cells, and spherical cysts. They can also form clusters of multiple cells. [1] [2]
In the flagellate swimming stage, the most common stage in their life cycle, the cells of Syssomonas resemble a typical opisthokont cell, reminiscent of animal sperm cells or chytrid zoospores. There is one smooth flagellum that emerges from the middle-lateral point of the cell, turns back, and directs backwards during swimming. While swimming, the fast beating of the flagellum can create the appearance of two flagella. The swimming cells rotate, and can suddenly stop and change direction of the movement. Solitary cells can attach temporarily to a substrate through the anterior part of the cell body, and produce water flow by rapid flagellar beating, resembling choanoflagellates or choanocytes from sponges. Floating cells move downwards to transform into amoeboflagellates by generating wide lobopodia and thin short filopodia, and slowing the flagellar beating. The amoeboflagellates can crawl along the substrate through their anterior lobopodia. [2]
The amoeboflagellate stages of Syssomonas can lose the flagellum by three different ways: discarding it abruptly, retracting it into the cell when stretched out, or coil under the cell and then retract into the cell as a spiral. As a result they become the amoeboid stage, which produces thin short filopodia and sometimes have two contractile vacuoles. Both amoebae and amoeboflagellates can turn back into flagellates. [2]
The amoeboid stage can retract its filopodia and become a round cyst, in which palintomic cell division (i.e. rapid cell divisions without cytoplasmic growth in between, a characteristic of animal embryonic cleavage) [3] can occur, generating 2, 4, 8 or 16 flagellated cells that are released from inside the cyst. [2]
Cells of Syssomonas can merge partially and form temporary aggregations of about 3–10 cells, usually shapeless and observed near the bottom of the water column. They can also aggregate by joining only flagellated cells together, with their flagella directed outwards, resembling the rosette-like colonies of choanoflagellates. Both aggregations break up easily, and their cell membranes are not fused. [2]
In solid cultures, solitary cells can sometimes merge completely at the bottom of the Petri dish into a syncytium-like or pseudoplasmodium structure, in which the nuclei do not merge. From these syncytia, budding of daughter cells occurs. This phenomenon of budding from syncytia has not been observed in any other eukaryotes, although the formation of multinucleated cells as a result of aggregation of multiple cells is known in other protist lineages (dictyostelids in Eumycetozoa, Copromyxa in Tubulinea, [4] acrasids in Excavata, [5] Sorogena in Alveolata, [6] Sorodiplophrys in Stramenopiles, Guttulinopsis in Rhizaria, [7] and Fonticula alba within the opisthokonts). [8] The transition from an amoeboid filopodial stage to an aggregative stage is also observed in another holozoan, Capsaspora owczarzaki . The formation of Syncytia also occurs in animals; the cytoplasm of glass sponges, teguments of flatworms, and the skeletal muscles and placenta of mammals are all syncytial structures. [2]
The merging of cells in Syssomonas attracts, likely by chemical signaling, other nearby cells that actively swim and try to attach to the aggregates. This appears to be the only method by which aggregates grow, as opposed to cell division. [2]
Syssomonas multiformis was isolated from a freshwater pool in Vietnam. The organism can survive temperatures ranging from 5 to 36 °C. It feeds on the cytoplasmic content of other eukaryotes of similar size, which is an unusual trait among unicellular holozoans. In particular, it is a predator of heterotrophic chrysomonads and bodonids (e.g. Parabodo caudatus and Spumella species). [2] They can also engulf bacteria and small debris, in a similar manner to choanoflagellates. [1]
In contrast to many other eukaryotic protists, Syssomonas cells do not possess any extruding organelles for hunting. Instead, they attach to the prey cell and suck out their cytoplasm without ingesting the cell membrane. They feed better on inactive, slow or dead cells or cysts. Likely by chemical signaling, after one cell attaches to the prey, many other Syssomonas cells become attracted to the same prey cell and try to attach to it. Several cells can suck out the cytoplasm of the same prey cell jointly. [2]
They use short pseudopodia to feed on clusters of bacteria. Afterwards, they form a large food vacuole at the posterior cell end. However, bacteria alone are not sufficient nutrition for Syssomonas: without any eukaryotic prey, their cells die or form resting cysts. [2]
As a lineage of Holozoa, Syssomonas is one of many protist groups closely related to animals and is therefore a subject of research in the search for the origin of animal multicellularity. The first phylogenomic analyses including Syssomonas recovered the genus as the sister taxon of Corallochytrium . Together, they compose the clade Pluriformea, which was recovered as the sister taxon of Filozoa. [1] An alternative hypothesis places Pluriformea as the sister group of Ichthyosporea in a clade known as Teretosporea. [2] The following cladogram displays the position of Syssomonas among the opisthokonts, according to the first hypothesis:
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.
A flagellum is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores (zoospores), and from a wide range of microorganisms to provide motility. Many protists with flagella are known as flagellates.
The choanoflagellates are a group of free-living unicellular and colonial flagellate eukaryotes considered to be the closest living relatives of the animals. Choanoflagellates are collared flagellates, having a funnel shaped collar of interconnected microvilli at the base of a flagellum. Choanoflagellates are capable of both asexual and sexual reproduction. They have a distinctive cell morphology characterized by an ovoid or spherical cell body 3–10 μm in diameter with a single apical flagellum surrounded by a collar of 30–40 microvilli. Movement of the flagellum creates water currents that can propel free-swimming choanoflagellates through the water column and trap bacteria and detritus against the collar of microvilli, where these foodstuffs are engulfed. This feeding provides a critical link within the global carbon cycle, linking trophic levels. In addition to their critical ecological roles, choanoflagellates are of particular interest to evolutionary biologists studying the origins of multicellularity in animals. As the closest living relatives of animals, choanoflagellates serve as a useful model for reconstructions of the last unicellular ancestor of animals. According to a 2021 study, crown group craspedids appeared 422.78 million years ago, Although a previous study from 2017 recovered the divergence of the crown group choanoflagellates (craspedids) at 786.62 million years.
The opisthokonts are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and Breviata comprise the larger clade Obazoa.
Naegleria is a genus consisting of 47 described species of protozoa often found in warm aquatic environments as well as soil habitats worldwide. It has three life cycle forms: the amoeboid stage, the cyst stage, and the flagellated stage, and has been routinely studied for its ease in change from amoeboid to flagellated stages. The Naegleria genera became famous when Naegleria fowleri, the causative agent of the usually fatal human and animal disease primary amoebic meningoencephalitis (PAM), was discovered in 1965. Most species in the genus, however, are incapable of causing disease.
Amorphea is a taxonomic supergroup that includes the basal Amoebozoa and Obazoa. That latter contains the Opisthokonta, which includes the Fungi, Animals and the Choanomonada, or Choanoflagellates. The taxonomic affinities of the members of this clade were originally described and proposed by Thomas Cavalier-Smith in 2002.
Capsaspora is a monotypic genus containing the single species Capsaspora owczarzaki. C. owczarzaki is a single-celled eukaryote that occupies a key phylogenetic position in our understanding of the origin of animal multicellularity, as one of the closest unicellular relatives to animals. It is, together with Ministeria vibrans, a member of the Filasterea clade. This amoeboid protist has been pivotal to unraveling the nature of the unicellular ancestor of animals, which has been proved to be much more complex than previously thought.
Holozoa is a clade of organisms that includes animals and their closest single-celled relatives, but excludes fungi and all other organisms. Together they amount to more than 1.5 million species of purely heterotrophic organisms, including around 300 unicellular species. It consists of various subgroups, namely Metazoa and the protists Choanoflagellata, Filasterea, Pluriformea and Ichthyosporea. Along with fungi and some other groups, Holozoa is part of the Opisthokonta, a supergroup of eukaryotes. Choanofila was previously used as the name for a group similar in composition to Holozoa, but its usage is discouraged now because it excludes animals and is therefore paraphyletic.
Mastigamoeba is a genus of pelobionts, and treated by some as members of the Archamoebae group of protists. Mastigamoeba are characterized as anaerobic, amitochondriate organisms that are polymorphic. Their dominant life cycle stage is as an amoeboid flagellate. Species are typically free living, though endobiotic species have been described.
The Filozoa are a monophyletic grouping within the Opisthokonta. They include animals and their nearest unicellular relatives.
An amoeba, often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopods. Amoebae do not form a single taxonomic group; instead, they are found in every major lineage of eukaryotic organisms. Amoeboid cells occur not only among the protozoa, but also in fungi, algae, and animals.
Choanozoa is a clade of opisthokont eukaryotes consisting of the choanoflagellates (Choanoflagellatea) and the animals. The sister-group relationship between the choanoflagellates and animals has important implications for the origin of the animals. The clade was identified in 2015 by Graham Budd and Sören Jensen, who used the name Apoikozoa. The 2018 revision of the classification first proposed by the International Society of Protistologists in 2012 recommends the use of the name Choanozoa.
Mesofila is a genus of freshwater heterotrophic protists of the phylum Cercozoa. It is the only genus in the family Mesofilidae. It is a monotypic genus, with the sole species M. limnetica.
The glissomonads are a group of bacterivorous gliding flagellated protists that compose the order Glissomonadida, in the amoeboflagellate phylum Cercozoa. They comprise a vast, largely undescribed diversity of soil and freshwater organisms. They are the sister group to cercomonads; the two orders form a solid clade of gliding soil-dwelling flagellates called Pediglissa.
Viridiraptoridae, previously known as clade X, is a clade of heterotrophic protists in the phylum Cercozoa. They're a family of glissomonads, a group containing a vast, mostly undescribed diversity of soil and freshwater organisms.
Helkesida is a group of microscopic protists belonging to the supergroup Rhizaria, both discovered through molecular phylogenetic analyses. It contains amoeboid flagellates with two flagella. They are either free-living, mostly on fecal matter, or live inside the gut of animals. Among these amoebae, one lineage has independently evolved aggregative multicellularity similarly to slime moulds.
Tunicaraptor is a genus of marine microbial protists containing the single species Tunicaraptor unikontum, discovered in 2020 from marine waters of Chile. It is a lineage of predatorial flagellates closely related to animals. It has a rare feeding structure not seen in other opisthokonts.
An amoeboflagellate is any eukaryotic organism capable of behaving as an amoeba and as a flagellate at some point during their life cycle. Amoeboflagellates present both pseudopodia and at least one flagellum, often simultaneously.
Naegleria lustrarea is a species of free-living amoeba belonging to the group Heterolobosea. It was described in 2024 from amoebae present in the gut of a salamander found in Arkansas. It is an amoeboflagellate, capable of transitioning from an amoeboid stage to a flagellate stage during its life cycle.
Idionectes is a genus of amoeba discovered from Allensbach, Konstanz, Germany. It contains only one species, I. vortex. Described by Sebastian Hess and Alastair G. B. Simpson in 2019, the scientific name means distinct or peculiar swimmer. Named because of its unique locomotion by creating a water vortex with its flying saucer-like body, it was also dubbed UFO. It is the only known eukaryotic cell having a rotating flagellum, which is the usual characteristic of prokaryotes.