Capsaspora | |
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Scientific classification | |
Domain: | Eukaryota |
Class: | Filasterea |
Order: | Ministeriida |
Family: | Capsasporidae Cavalier-Smith, 2008 [1] |
Genus: | Capsaspora Hertel, Bayne & Loker 2002 [2] |
Species: | C. owczarzaki |
Binomial name | |
Capsaspora owczarzaki Hertel, Bayne & Loker 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 (see “Taxonomy” below). 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. [3] [4] [5]
C. owczarzaki was originally described as an amoeba-like “symbiont” of the fresh-water snail Biomphalaria glabrata . [6] [7] The amoebae were obtained from the haemolymph of snails originally sampled in Puerto Rico.
C. owczarzaki’s life cycle comprises 3 different stages with three different cell types, which was reported in 2017. [8] Under culture conditions, C. owczarzaki’s filopodial cells crawl attached to the substrate, with active replication until the end of the exponential growth phase. Then, cells start to detach, retracting the branching filopodia and encysting. During this cystic phase, division is stopped. Alternatively, amoebae can actively aggregate to each other by unknown factors, forming a multicellular, aggregative structure and secreting an unstructured extracellular material that seems to prevent direct cell-to-cell contact.
C. owczarzaki cells, in the filopodial stage, were described as 3 to 5 μm amoebas with a nucleus ⅓ - ½ of the diameter of the cell (containing a central nucleolus), long branched filopodia, mitochondria with flattened cristae, numerous phagosomes, lipid vacuoles, glycogen granules and a Golgi apparatus. [6] [7] Cystic cells measure 4 to 5 μm with a double wall: the outer thin, irregular and loosely attached; and the inner thicker, smooth. [9]
C. owczarzaki is together with Ministeria vibrans a member of the Filasterea clade. [10] [1] This group is the sister group to a clade comprising Metazoa and Choanoflagellata, which together form the Filozoa [10] [1] [11] (See Figure 1). C. owczarzaki was originally described as nucleariids. [7] However, later molecular ribosomal phylogenies placed C. owczarzaki somewhere closer to animals than the rest of nucleariids. [12] [13] [14] Finally, a multi-gene phylogenetic analysis with several opisthokont taxa clearly showed that C. owczarzaki is not a nucleariid, but part of the Holozoa. [15] [16] This was later on corroborated by phylogenomic analyses, [17] [18] one of which [18] situated it as sister-group to Ministeria forming the Filasterea clade, which is the sister-group to Choanoflagellatea and Metazoa. [19] [20]
C. owczarzaki is of scientific interest because it is one of the closest unicellular relatives of multicellular animals. Its genome has recently been sequenced and shows several genes involved in metazoan multicellularity, such as integrins, [21] metazoan transcription factors, [22] and protein tyrosine kinases. [23] Moreover, it has relevance to human health because its host, the snail Biomphalaria glabrata , is also the intermediate host of the digenean flatworm Schistosoma mansoni , the causative agent of widespread schistosomiasis in humans. C. owczarzaki not only parasitizes the intermediate host of S. mansoni but also attacks and kills the sporocysts of the flatworm living inside the snail. [24]
Nucleariida is a group of amoebae with filose pseudopods, known mostly from soils and freshwater. They are distinguished from the superficially similar vampyrellids mainly by having mitochondria with discoid cristae, in the absence of superficial granules, and in the way they consume food.
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.
The Rhizaria are a diverse and species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthetic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has been described. This group was used by Cavalier-Smith in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon.
Cristidiscoidea or Nucleariae is a proposed basal holomycota clade in which Fonticula and Nucleariida emerged, as sister of the fungi. Since it is close to the divergence between the main lineages of fungi and animals, the study of Cristidiscoidea can provide crucial information on the divergent lifestyles of these groups and the evolution of opisthokonts and slime mold multicellularity. The holomycota tree is following Tedersoo et al.
The Apusozoa are a paraphyletic phylum of flagellate eukaryotes. They are usually around 5–20 μm in size, and occur in soils and aquatic habitats, where they feed on bacteria. They are grouped together based on the presence of an organic shell or theca under the dorsal surface of the cell.
The Urmetazoan is the hypothetical last common ancestor of all animals, or metazoans. It is universally accepted to be a multicellular heterotroph — with the novelties of a germline and oogamy, an extracellular matrix (ECM) and basement membrane, cell-cell and cell-ECM adhesions and signaling pathways, collagen IV and fibrillar collagen, different cell types, spatial regulation and a complex developmental plan, and relegated unicellular stages.
Ministeria vibrans is a bacterivorous amoeba with filopodia that was originally described to be suspended by a flagellum-like stalk attached to the substrate. Molecular and experimental work later on demonstrated the stalk is indeed a flagellar apparatus.
Breviata anathema is a single-celled flagellate amoeboid eukaryote, previously studied under the name Mastigamoeba invertens. The cell lacks mitochondria, much like the pelobionts to which the species was previously assigned, but has remnant mitochondrial genes, and possesses an organelle believed to be a modified anaerobic mitochondrion, similar to the mitosomes and hydrogenosomes found in other eukaryotes that live in low-oxygen environments.
Filasterea is a proposed basal Filozoan clade of single-celled ameboid eukaryotes that includes Ministeria and Capsaspora. It is a sister clade to the Choanozoa in which the Choanoflagellatea and Animals appeared, originally proposed by Shalchian-Tabrizi et al. in 2008, based on a phylogenomic analysis with 78 genes. Filasterea was found to be the sister-group to the clade composed of Metazoa and Choanoflagellata within the Opisthokonta, a finding that has been further corroborated with additional, more taxon-rich, phylogenetic analyses.
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.
The apusomonads are a group of protozoan zooflagellates that glide on surfaces, and mostly consume prokaryotes. They are of particular evolutionary interest because they appear to be the sister group to the Opisthokonts, the clade that includes both animals and fungi. Together with the Breviatea, these form the Obazoa clade.
The Filozoa are a monophyletic grouping within the Opisthokonta. They include animals and their nearest unicellular relatives.
Holomycota or Nucletmycea are a basal Opisthokont clade as sister of the Holozoa. It consists of the Cristidiscoidea and the kingdom Fungi. The position of nucleariids, unicellular free-living phagotrophic amoebae, as the earliest lineage of Holomycota suggests that animals and fungi independently acquired complex multicellularity from a common unicellular ancestor and that the osmotrophic lifestyle was originated later in the divergence of this eukaryotic lineage. Opisthosporidians is a recently proposed taxonomic group that includes aphelids, Microsporidia and Cryptomycota, three groups of endoparasites.
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.
Creolimax fragrantissima is a single-celled protist that occupies a key phylogenetic position to understand the origin of animals. It was isolated from the digestive tract of some marine invertebrates, mainly from the peanut worm, collected from the Northeast Pacific.
Sphaeroforma arctica, is a unicellular eukaryote with a pivotal position in the tree of life. It was first isolated from the arctic marine amphipod Gammarus setosus. Like other Ichthyosporeans such as Creolimax and Abeoforma, Sphaeroforma arctica are spherical cells characterized with their capacity to grow into multi-nucleated coenocytes. However, a unique feature of S. arctica, is that no obvious budding, hyphal, amoeboid, sporal or flagellated growth stages have been observed in laboratory growth conditions.
Mantamonads are a group of free-living heterotrophic flagellates that move primarily by gliding on surfaces. They are classified as one genus Mantamonas in the monotypic family Mantamonadidae, order Mantamonadida and class Glissodiscea. Previously, they were classified in Apusozoa as sister of the Apusomonadida on the basis of rRNA analyses. However, mantamonads are currently placed in CRuMs on the basis of phylogenomic analyses that identify their closest relatives as the Diphylleida and Rigifilida.
Parvularia atlantis is a filopodiated amoeba which was isolated from a lake in Atlanta and deposited in the American Type Culture Collection (ATCC) under the name Nuclearia sp. ATCC 50694 on 1997 by TK Sawyer. It was classified under the genus Nuclearia and morphologically resembles to Nuclearia species, although it is smaller. Later it was determined that it phylogenetically belongs to a new nucleariid lineage., distantly related to Nuclearia and Fonticula genera – the other two previously described nucleariid genera.
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.
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.