Syssomonas

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Syssomonas
Syssomonas 2X 2020.webp
SEM image of Syssomonas.
ac = acroneme, fl = flagellum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
(unranked): Opisthokonta
(unranked): Holozoa
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 [lower-alpha 1]
Species
  • S. multiformis

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.

Contents

Life stages

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]

Unicellular stages: flagellar, amoeboid and cyst

External morphology and life forms of Syssomonas multiformis. A-B-C: swimming flagellated cells. D: amoeboflagellate (lb = lobopodia). E-F: amoeboid cells (fp = filopodia). G: cyst. H: palintomic cell division inside a cyst. I: cyst with vesicles. J: attached flagellated cell. Syssomonas life forms.png
External morphology and life forms of Syssomonas multiformis. A-B-C: swimming flagellated cells. D: amoeboflagellate (lb = lobopodia). E-F: amoeboid cells (fp = filopodia). G: cyst. H: palintomic cell division inside a cyst. I: cyst with vesicles. J: attached flagellated cell.

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]

Aggregative multicellular stages

K-M, O: cellular aggregations of Syssomonas near the bottom of the Petri dish. N: floating aggregate of flagellated cells. Syssomonas aggregates.webp
K-M, O: cellular aggregations of Syssomonas near the bottom of the Petri dish. N: floating aggregate of flagellated cells.

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]

Ecology

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]

Evolution

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:

Opisthokonta
Holozoa
Filozoa
Choanozoa

Metazoa Cetoine doree vol.jpg

Choanoflagellata Monosiga Brevicollis Phase.jpg

Filasterea

Pigoraptor

Capsaspora owczarzaki Capsaspora owczarzaki.jpeg

Ministeria vibrans Ministeria vibrans.jpeg

Pluriformea

Syssomonas multiformis Syssomonas 2X 2020.webp

Corallochytrium limacisporum Corallochytrium limacisporum.png

Ichthyosporea Sphaeroforma arctica.jpg

Holomycota Linear arrangement of ascospores in the asci of the fungus Sordaria macrospora Cropped.jpg

Notes

  1. Deposited in the Marine Invertebrate Collection, Beaty Biodiversity Museum, University of British Columbia

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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.

<span class="mw-page-title-main">Flagellum</span> Cellular appendage functioning as locomotive or sensory organelle

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<span class="mw-page-title-main">Choanoflagellate</span> Group of eukaryotes considered the closest living relatives of animals

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.

<span class="mw-page-title-main">Opisthokont</span> Group of eukaryotes which includes animals and fungi, among other groups

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<span class="mw-page-title-main">Amorphea</span> Members of the Unikonta, a taxonomic group proposed by Thomas Cavalier-Smith

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{{Paraphyletic group | classification_status = paraphyletic | name = Protists | fossil range = {{Long fossil range|2100|0|[[Paleopr | image = Protist collage 2.jpg | image_alt = | image_caption = Examples of protists. Clockwise from top left: red algae, kelp, ciliate, golden alga, dinoflagellate, metamonad, amoeba, slime mold. | auto = yes | parent = Eukaryota | subdivision_ranks = Supergroups |subdivision= Amorphea (including fungi & animals)
Apusomonadida
Archaeplastida (including land plants)
Breviatea
CRuMs
Cryptista
Discoba
Haptista
Hemimastigophora
Malawimonadida
Metamonada
Provora
SAR supergroup
Telonemia | excludes = Animalia
Fungi
Embryophyta
}}

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