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The protosteloid amoebae, or protosteloids (formerly known as protostelids), are a group of terrestrial
amoebae An amoeba (; less commonly spelled ameba or amœba; : amoebas (less commonly, amebas) or amoebae (amebae) ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and r ...
capable of developing a tiny fruiting body or sporocarp consisting of a stalk supporting one or more
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s. They do not form a natural group; instead, they have gained the sporocarp-forming ability independently from each other during evolution.


Description

Protosteloid amoebae are capable of making simple fruiting bodies consisting of a cellular stalk topped by one or a few
spores In biology, a spore is a unit of sexual (in fungi) or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plant ...
. These are known as sporocarps. All species are
microscopic The microscopic scale () is the scale of objects and events smaller than those that can easily be seen by the naked eye, requiring a lens or microscope to see them clearly. In physics, the microscopic scale is sometimes regarded as the scale betwe ...
and are typically found on dead plant matter where they consume
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
,
yeasts Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constitu ...
, and
fungal A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as one of the tradit ...
spores. Since protostelids are amoebae that make spores, they are considered to be
slime molds Slime mold or slime mould is an informal name given to a polyphyly, polyphyletic assemblage of unrelated eukaryotic organisms in the Stramenopiles, Rhizaria, Discoba, Amoebozoa and Holomycota clades. Most are near-microscopic; those in the Myx ...
. Protosteloid amoebae are terrestrial, typically found on dead plant matter, including stems and
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
of
herbaceous plants Herbaceous plants are vascular plants that have no persistent wood, woody stems above ground. This broad category of plants includes many perennial plant, perennials, and nearly all Annual plant, annuals and Biennial plant, biennials. Definition ...
, stems and leaves of
grasses Poaceae ( ), also called Gramineae ( ), is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos, the grasses of natural grassland and species cultivated in ...
,
bark Bark may refer to: Common meanings * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Arts and entertainment * ''Bark'' (Jefferson Airplane album), ...
of living trees, decaying wood and other types of dead plant matter. Some species are aquatic; they live on dead plant parts submerged in a pond. Others live on the petals of living flowers and on living tree leaves. Since protosteloid amoebae eat bacteria, yeasts, and fungal spores in the laboratory, it is thought that they also do this in nature. They are thus thought to be predators in
decomposer Decomposers are organisms that break down dead organisms and release the nutrients from the dead matter into the environment around them. Decomposition relies on chemical processes similar to digestion in animals; in fact, many sources use the word ...
communities.


Diversity

Protosteloid development has evolved numerous times in different
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s of
Amoebozoa Amoebozoa is a major Taxonomy (biology), taxonomic group containing about 2,400 described species of Amoeba, amoeboid protists, often possessing blunt, fingerlike, Pseudopod#Morphology, lobose pseudopods and tubular mitochondrial cristae. In trad ...
, a major clade of mostly
amoeboid An amoeba (; less commonly spelled ameba or amœba; : amoebas (less commonly, amebas) or amoebae (amebae) ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and r ...
eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
s. Three of the major groups of amoebozoans,
Discosea Discosea is a class of Amoebozoa, consisting of naked amoebae with a flattened, discoid body shape. Members of the group do not produce tubular or subcylindrical pseudopodia, like amoebae of the class Tubulinea. When a discosean is in motion, a ...
, Variosea and
Eumycetozoa Eumycetozoa (), or true slime molds, is a diverse group of protists that behave as slime molds and develop fruiting bodies, either as sorocarps or as sporocarp (fungi), sporocarps. It is a monophyletic group or clade within the phylum Amoebozoa t ...
, have at least some protosteloid members, with Variosea being the more abundant in protosteloids. They are entirely absent in Tubulinea, the fourth major group. Within Variosea, Cavosteliida is the largest protosteloid group, containing four genera: '' Cavostelium'', '' Nannostelium'', '' Schizoplasmodiopsis'' and '' Tychosporium''. Another group is Protosteliida, containing the genus '' Protostelium'', the first protosteloid organism to be discovered. In Eumycetozoa are the protosteloid genera '' Protosporangium'' within Protosporangiida, and '' Echinostelium'' and '' Echinosteliopsis'' within
Myxogastria Myxogastria/Myxogastrea (myxogastrids, ICZN) or Myxomycetes ( ICN) is a class of slime molds that contains 5  orders, 14  families, 62 genera, and 888 species. They are colloquially known as the ''plasmodial'' or ''acellula ...
. Within Discosea, several orders contain amoebae that have evolved protosteloid development: the genus '' Luapeleamoeba'' and the species '' Acanthamoeba pyriformis'' in the order Centramoebida; the genus '' Endostelium'' in the order Pellitida; the genus '' Protosteliopsis'' in the order Vannellida; and the genus '' Microglomus'' in the order Dermamoebida.


Distribution

Collections of protosteloids have been made from all continents, including the
Antarctic peninsula The Antarctic Peninsula, known as O'Higgins Land in Chile and Tierra de San Martin in Argentina, and originally as Graham Land in the United Kingdom and the Palmer Peninsula in the United States, is the northernmost part of mainland Antarctica. ...
. Protosteloids have also been found on isolated islands like
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
in the
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is bounded by the cont ...
and
Ascension Island Ascension Island is an isolated volcanic island, 7°56′ south of the Equator in the Atlantic Ocean, South Atlantic Ocean. It is about from the coast of Africa and from the coast of South America. It is governed as part of the British Overs ...
in the southern
Atlantic The Atlantic Ocean is the second largest of the world's five oceanic divisions, with an area of about . It covers approximately 17% of Earth's surface and about 24% of its water surface area. During the Age of Discovery, it was known for se ...
, indicating that protosteloids have a worldwide distribution. Most studies of protosteloid distribution have been done in the
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
zones so they are best known from these areas. However,
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
studies have turned up protosteloids, often in great abundance.


Collection and laboratory culture

Since protosteloid amoebae are microscopic one must bring their substrates, dead plant matter, into the laboratory to find them. Dead plant matter is placed on the
agar Agar ( or ), or agar-agar, is a jelly-like substance consisting of polysaccharides obtained from the cell walls of some species of red algae, primarily from " ogonori" and " tengusa". As found in nature, agar is a mixture of two components, t ...
surface in a petri plate and allowed to incubate for several days to a week. Then the edges of the substrates are scanned with a compound microscope and species are identified by their fruiting body
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
and amoebal morphology. When protosteloid fruiting bodies are found they can be moved into laboratory
culture Culture ( ) is a concept that encompasses the social behavior, institutions, and Social norm, norms found in human societies, as well as the knowledge, beliefs, arts, laws, Social norm, customs, capabilities, Attitude (psychology), attitudes ...
onto an appropriate food organism or mix of organisms. This is done by picking up fruiting bodies or spores with a sterilised needle and moving them onto agar in a fresh petri plate that has been smeared with a bacterium or yeast upon which the protosteloid amoeba species has been known to grow. If the spores
germinate Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an flowering plant, angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the sp ...
then the protostelid begins eating the food organism and a culture is established.


Notes


References

{{Amoebozoa Amoeboids