Sphaeroforma arctica | |
---|---|
Sphaeroforma arctica stained with phalloidin and DAPI | |
Scientific classification | |
Domain: | Eukaryota |
(unranked): | Holozoa |
Class: | Ichthyosporea |
Order: | Ichthyophonida |
Family: | Creolimacidae |
Genus: | Sphaeroforma |
Species: | S. arctica |
Binomial name | |
Sphaeroforma arctica | |
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 . [2] [1] Like other Ichthyosporeans such as Creolimax and Abeoforma, Sphaeroforma arctica are spherical cells characterized with their capacity to grow into multi-nucleated coenocytes (multi-nucleates cell). 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.
Sphaeroforma is a member of the Ichthyosporea clade, which is the earliest branching holozoan lineage. It is a key organism to understand the origin of animals.
Sphaeroforma arctica grows easily in marine broth, with the particularity of growing synchronously. [3] [4] From a cell with 1 single nucleus (although newborn cells have been shown to contain 2 or even 4 nuclei, too [4] ), it can grow to reach a 128 nuclei cell before undergoing cellularization (the process by which the coenocytes split up to give new-born cells). Cellularization involves coordinated inward plasma membrane invaginations dependent on an actomyosin network that leads to the formation of a polarized layer of cells resembling an epithelium. This process is associated with tightly regulated transcriptional activation of genes involved in cell adhesion. [5]
Moreover, it has been recently shown that S. arctica contains miRNA as well as the complex miRNA processing machinery. Taken together, Sphaeroforma arctica is a great model organism from both evolutionary and cell biology perspectives. [6]
The cell nucleus is a membrane-bound organelle found in eukaryotic cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, have no nuclei, and a few others including osteoclasts have many. The main structures making up the nucleus are the nuclear envelope, a double membrane that encloses the entire organelle and isolates its contents from the cellular cytoplasm; and the nuclear matrix, a network within the nucleus that adds mechanical support.
Mitosis is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis is an equational division which gives rise to genetically identical cells in which the total number of chromosomes is maintained. Mitosis is preceded by the S phase of interphase and is followed by telophase and cytokinesis, which divide the cytoplasm, organelles, and cell membrane of one cell into two new cells containing roughly equal shares of these cellular components. The different stages of mitosis altogether define the mitotic phase of a cell cycle—the division of the mother cell into two daughter cells genetically identical to each other.
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.
A coenocyte is a multinucleate cell which can result from multiple nuclear divisions without their accompanying cytokinesis, in contrast to a syncytium, which results from cellular aggregation followed by dissolution of the cell membranes inside the mass. The word syncytium in animal embryology is used to refer to the coenocytic blastoderm of invertebrates. A coenocytic colony is referred to as a coenobium, and most coenobia are composed of a distinct number of cells, often as a multiple of two.
Acoelomorpha is a subphylum of very simple and small soft-bodied animals with planula-like features which live in marine or brackish waters. They usually live between grains of sediment, swimming as plankton, or crawling on other organisms, such as algae and corals. With the exception of two acoel freshwater species, all known acoelomorphs are marine.
In evolutionary biology, the term cellularization (cellularisation) has been used in theories to explain the evolution of cells, for instance in the pre-cell theory, dealing with the evolution of the first cells on this planet, and in the syncytial theory attempting to explain the origin of Metazoa from unicellular organisms.
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 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.
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 unravel the nature of the unicellular ancestor of animals, which has been proved to be much more complex than previously thought.
Corallochytrium belongs to the class of Corallochytrea within Teretosporea and is a sister group to Ichthyosporea. Corallochytrium limacisporum is the only species of Corallochytrium known so far. It was first discovered and named in the Arabian Sea’s coral lagoons by Kaghu-Kumar in 1987. It was first thought to be a member of the fungi-like thraustochytrids, however, this was later disproven due to Corallochytriums lack of cilia and sagenogenetosome. Little research has been done on the life cycle or morphology. Most research concerning this genus has been done to uncover the evolution of animals and fungi, as Corallochytrium possess both animal and fungal enzymatic trademarks.
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 dozens of 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 eukaryotes constitute the domain of Eukarya or Eukaryota, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, and many unicellular organisms are eukaryotes. They constitute a major group of life forms alongside the two groups of prokaryotes: the Bacteria and the Archaea. Eukaryotes represent a small minority of the number of organisms, but given their generally much larger size, their collective global biomass is much larger than that of prokaryotes.
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.
Until the late 1950s, the Precambrian was not believed to have hosted multicellular organisms. However, with radiometric dating techniques, it has been found that fossils initially found in the Ediacara Hills in Southern Australia date back to the late Precambrian. These fossils are body impressions of organisms shaped like disks, fronds and some with ribbon patterns that were most likely tentacles.
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.
Abeoforma whisleri is a single-celled eukaryote that belongs to the Ichthyosporea clade, a group of protists closely related to animals.
Ichthyophonus hoferi is a single-celled protist that occupies a key phylogenetic position to understand the origin of animals. It has chitin cell wall, hyphae, and an amoeboid stage. It is a common parasite of marine and freshwater fishes.
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 nucleriid genera.
Chromosphaera perkinsii is a species of Ichthyosporea from the order Dermocystida. Named after Professor Frank Perkins, it was isolated in shallow marine sediments in Hawaii by Stuart Donachie and collaborators. It is a rare case of a putatively free-living ichthyosporean, and possibly the only free-living dermocystid.