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The Radiolaria, also called Radiozoa, are
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
of diameter 0.1–0.2 mm that produce intricate
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. ( ...
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
s, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ectoplasm. The elaborate mineral skeleton is usually made of
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
. They are found as
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
throughout the global ocean. As zooplankton, radiolarians are primarily
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic, but many have photosynthetic endosymbionts and are, therefore, considered mixotrophs. The skeletal remains of some types of radiolarians make up a large part of the cover of the ocean floor as
siliceous ooze Siliceous ooze is a type of biogenic pelagic sediment located on the deep ocean floor. Siliceous oozes are the least common of the deep sea sediments, and make up approximately 15% of the ocean floor. Oozes are defined as sediments which contain a ...
. Due to their rapid change as species and intricate skeletons, radiolarians represent an important diagnostic
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
found from the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
onwards.


Description

Radiolarians have many needle-like pseudopods supported by bundles of
microtubule Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
s, which aid in the radiolarian's buoyancy. The
cell nucleus The cell nucleus (pl. nuclei; from Latin or , meaning ''kernel'' or ''seed'') 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, h ...
and most other
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s are in the endoplasm, while the ectoplasm is filled with frothy
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s and
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
droplets, keeping them buoyant. The radiolarian can often contain
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
algae, especially zooxanthellae, which provide most of the cell's energy. Some of this organization is found among the
heliozoa Heliozoa, commonly known as sun-animalcules, are microbial eukaryotes (protists) with stiff arms ( axopodia) radiating from their spherical bodies, which are responsible for their common name. The axopodia are microtubule-supported projections fro ...
, but those lack central capsules and only produce simple scales and spines. Some radiolarians are known for their resemblance to regular polyhedra, such as the
icosahedron In geometry, an icosahedron ( or ) is a polyhedron with 20 faces. The name comes and . The plural can be either "icosahedra" () or "icosahedrons". There are infinitely many non- similar shapes of icosahedra, some of them being more symmetrica ...
-shaped ''Circogonia icosahedra'' pictured below.


Taxonomy

The radiolarians belong to the supergroup Rhizaria together with ( amoeboid or
flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
)
Cercozoa Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead defined by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eu ...
and (shelled amoeboid)
Foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
. Traditionally the radiolarians have been divided into four groups— Acantharea,
Nassellaria Nassellaria is an order of Rhizaria belonging to the class Radiolaria. The organisms of this order are characterized by a skeleton cross link with a cone or ring. Introduction Nassellaria is an order of Radiolaria under the class Polycystina. ...
,
Spumellaria Spumellaria is an order of radiolarians in the class Polycystinea. They are ameboid protists appearing in abundance in the world's oceans, possessing a radially-symmetrical silica (opal) skeleton that has ensured their preservation in fossil rec ...
and
Phaeodarea Phaeodarea, or Phaeodaria, is a group of amoeboid cercozoan organisms. They are traditionally considered radiolarians, but in molecular trees do not appear to be close relatives of the other groups, and are instead placed among the Cercozoa. They ...
. Phaeodaria is however now considered to be a Cercozoan. Nassellaria and Spumellaria both produce siliceous skeletons and were therefore grouped together in the group
Polycystina The polycystines are a group of radiolarians. They include the vast majority of the fossil radiolaria, as their skeletons are abundant in marine sediments, making them one of the most common groups of microfossils. These skeletons are composed ...
. Despite some initial suggestions to the contrary, this is also supported by molecular phylogenies. The Acantharea produce skeletons of
strontium sulfate Strontium sulfate (SrSO4) is the sulfuric acid, sulfate salt of strontium. It is a white crystalline powder and occurs in nature as the mineral Celestine (mineral), celestine. It is poorly soluble in water to the extent of 1 part in 8,800. It is mo ...
and is closely related to a peculiar genus, ''
Sticholonche ''Sticholonche'' is a genus of radiolarians with a single species, ''Sticholonche zanclea'', found in open oceans at depths of 99–510 metres. It is generally considered a heliozoan, placed in its own order, called the Taxopodida. However it h ...
'' (
Taxopodida ''Sticholonche'' is a genus of radiolarians with a single species, ''Sticholonche zanclea'', found in open oceans at depths of 99–510 metres. It is generally considered a heliozoan, placed in its own order, called the Taxopodida. However it h ...
), which lacks an internal skeleton and was for long time considered a
heliozoa Heliozoa, commonly known as sun-animalcules, are microbial eukaryotes (protists) with stiff arms ( axopodia) radiating from their spherical bodies, which are responsible for their common name. The axopodia are microtubule-supported projections fro ...
n. The Radiolaria can therefore be divided into two major lineages: Polycystina (Spumellaria + Nassellaria) and Spasmaria (Acantharia + Taxopodida). There are several higher-order groups that have been detected in molecular analyses of environmental data. Particularly, groups related to Acantharia and Spumellaria. These groups are so far completely unknown in terms of morphology and physiology and the radiolarian diversity is therefore likely to be much higher than what is currently known. The relationship between the Foraminifera and Radiolaria is also debated. Molecular trees supports their close relationship—a grouping termed Retaria. But whether they are sister lineages or if the Foraminifera should be included within the Radiolaria is not known.


Biogeography

In the diagram on the right, a Illustrates generalized radiolarian provinces and their relationship to water mass temperature (warm versus cool color shading) and circulation (gray arrows). Due to high-latitude water mass submergence under warm, stratified waters in lower latitudes, radiolarian species occupy habitats at multiple latitudes, and depths throughout the world oceans. Thus, marine sediments from the tropics reflect a composite of several vertically stacked faunal assemblages, some of which are contiguous with higher latitude surface assemblages. Sediments beneath polar waters include cosmopolitan deep-water radiolarians, as well as high-latitude endemic surface water species. Stars in (a) indicate the latitudes sampled, and the gray bars highlight the radiolarian assemblages included in each sedimentary composite. The horizontal purple bars indicate latitudes known for good radiolarian (silica) preservation, based on surface sediment composition. Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
Data show that some species were extirpated from high latitudes but persisted in the tropics during the late
Neogene The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
, either by migration or range restriction (b). With predicted global warming, modern
Southern Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-small ...
species will not be able to use migration or range contraction to escape environmental stressors, because their preferred cold-water habitats are disappearing from the globe (c). However, tropical endemic species may expand their ranges toward midlatitudes. The color polygons in all three panels represent generalized radiolarian biogeographic provinces, as well as their relative water mass temperatures (cooler colors indicate cooler temperatures, and vice versa). File:Circogoniaicosahedra ekw.jpg, ''Circogonia icosahedra'', radiolarian species shaped like a regular icosahedron File:Anthocyrtium hispidum Haeckel - Radiolarian (34986365113).jpg, ''Anthocyrtium hispidum Haeckel''


Radiolarian shells

Radiolarians are unicellular predatory
protists A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the excl ...
encased in elaborate globular shells usually made of silica and pierced with holes. Their name comes from the Latin for "radius". They catch prey by extending parts of their body through the holes. As with the silica frustules of diatoms, radiolarian shells can sink to the ocean floor when radiolarians die and become preserved as part of the ocean sediment. These remains, as microfossils, provide valuable information about past oceanic conditions.Wassilieff, Maggy (2006
"Plankton - Animal plankton"
''Te Ara - the Encyclopedia of New Zealand''. Accessed: 2 November 2019.
File:Mikrofoto.de-Radiolarien 6.jpg, Like diatoms, radiolarians come in many shapes File:Theocotylissa ficus Ehrenberg - Radiolarian (34638920262).jpg, Also like diatoms, radiolarian shells are usually made of silicate File:Acantharian radiolarian Xiphacantha (Haeckel).jpg, However acantharian radiolarians have shells made from
strontium sulfate Strontium sulfate (SrSO4) is the sulfuric acid, sulfate salt of strontium. It is a white crystalline powder and occurs in nature as the mineral Celestine (mineral), celestine. It is poorly soluble in water to the extent of 1 part in 8,800. It is mo ...
crystals
File:Spherical radiolarian 2.jpg, Cutaway schematic diagram of a spherical radiolarian shell File:Cladococcus abietinus.jpg, ''Cladococcus abietinus''


Fossil record

The earliest known radiolaria date to the very start of the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
period, appearing in the same beds as the first small shelly fauna—they may even be terminal Precambrian in age. They have significant differences from later radiolaria, with a different silica lattice structure and few, if any, spikes on the
test Test(s), testing, or TEST may refer to: * Test (assessment), an educational assessment intended to measure the respondents' knowledge or other abilities Arts and entertainment * ''Test'' (2013 film), an American film * ''Test'' (2014 film), ...
. About ninety percent of known radiolarian species are extinct. The skeletons, or tests, of ancient radiolarians are used in
geological dating Geochronology is the science of determining the age of rocks, fossils, and sediments using signatures inherent in the rocks themselves. Absolute geochronology can be accomplished through radioactive isotopes, whereas relative geochronology is pr ...
, including for oil exploration and determination of ancient climates. Some common radiolarian fossils include '' Actinomma'', '' Heliosphaera'' and '' Hexadoridium''.


See also

* Radiolarite


References

* * * *


External links

*Radiolarians
*
Radiolaria.org
*
Tree Of Life—Radiolaria
{{Authority control * Amoeboids Extant Cambrian first appearances Bikont subphyla Taxa named by Thomas Cavalier-Smith