Protist Skeletons
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Many protists have protective shells or
tests 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) ...
, usually made from
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , commonly found in nature as quartz. In many parts of the world, silica is the major constituent of sand. Silica is one of the most complex and abundant f ...
(glass) or
calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
(chalk).
Protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...
s are a diverse group of
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 ...
organisms that are not plants, animals, or fungi. They are typically microscopic
unicellular organism A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
s that live in water or moist environments. Protists shells are often tough, mineralised forms that resist degradation, and can survive the death of the protist as a
microfossil A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
. Although protists are typically very small, they are ubiquitous. Their numbers are such that their shells play a huge part in the formation of
ocean sediment Marine sediment, or ocean sediment, or seafloor sediment, are deposits of insoluble particles that have accumulated on the seafloor. These particles either have their origins in soil and rocks and have been transported from the land to the ...
s and in the global cycling of elements and nutrients. The role of protist shells depends on the type of protist. Protists such as
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s and
radiolaria The Radiolaria, also called Radiozoa, are unicellular eukaryotes of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ect ...
have intricate, glass-like shells made of
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , commonly found in nature as quartz. In many parts of the world, silica is the major constituent of sand. Silica is one of the most complex and abundant f ...
that are hard and protective, and serve as a barrier to prevent water loss. The shells have small pores that allow for gas exchange and nutrient uptake.
Coccolithophore Coccolithophores, or coccolithophorids, are single-celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdom ...
s and
foraminifera Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
also have hard protective shells, but the shells are made of
calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
. These shells can help with buoyancy, allowing the organisms to float in the
water column The (oceanic) water column is a concept used in oceanography to describe the physical (temperature, salinity, light penetration) and chemical ( pH, dissolved oxygen, nutrient salts) characteristics of seawater at different depths for a defined ...
and move around more easily. In addition to protection and support, protist shells also serve scientists as a means of identification. By examining the characteristics of the shells, different species of protists can be identified and their ecology and evolution can be studied.


Protists

Cellular life likely originated as single-celled prokaryotes (including modern bacteria and archaea) and later evolved into more complex eukaryotes. Eukaryotes include organisms such as plants, animals, fungi and "protists". Protists are usually single-celled and microscopic. They can be
heterotrophic 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 ...
, meaning they obtain nutrients by consuming other organisms, or
autotrophic An autotroph is an organism that can convert abiotic sources of energy into energy stored in organic compounds, which can be used by other organisms. Autotrophs produce complex organic compounds (such as carbohydrates, fats, and proteins) us ...
, meaning they produce their own food through
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
or
chemosynthesis In biochemistry, chemosynthesis is the biological conversion of one or more carbon-containing molecules (usually carbon dioxide or methane) and nutrients into organic matter using the oxidation of inorganic compounds (e.g., hydrogen gas, hydrog ...
, or
mixotrophic A mixotroph is an organism that uses a mix of different sources of energy and carbon, instead of having a single trophic mode, on the continuum from complete autotrophy to complete heterotrophy. It is estimated that mixotrophs comprise more than ...
, meaning they produce their own food through a mixture of those methods. The term protist came into use historically to refer to a group of biologically similar organisms; however, modern research has shown it to be a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
group that does not contain all descendants of a common ancestor. As such it does not constitute a
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 ...
and is not currently in formal scientific use. Nonetheless, the term continues to be used informally to refer to those
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 that cannot be classified as plants, fungi or animals. Most protists are too small to be seen with the naked eye. They are highly diverse organisms currently organised into 18 phyla, but are not easy to classify. Studies have shown high protist diversity exists in oceans, deep sea-vents and river sediments, suggesting large numbers of eukaryotic microbial communities have yet to be discovered. As eukaryotes, protists possess within their cell at least one
nucleus Nucleus (: nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucleu ...
, as well as
organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as Organ (anatomy), organs are to th ...
s such as
mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...
and
Golgi bodies The Golgi apparatus (), also known as the Golgi complex, Golgi body, or simply the Golgi, is an organelle found in most eukaryotic Cell (biology), cells. Part of the endomembrane system in the cytoplasm, it protein targeting, packages proteins ...
. Many protists are asexual but can reproduce rapidly through
mitosis Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...
or by fragmentation; others (including
foraminifera Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
) may reproduce either sexually or asexually. In contrast to the cells of bacteria and archaea, the cells of protists and other eukaryotes are highly organised. Plants, animals and fungi are usually multi-celled and are typically
macroscopic The macroscopic scale is the length scale on which objects or phenomena are large enough to be visible with the naked eye, without magnifying optical instruments. It is the opposite of microscopic. Overview When applied to physical phenome ...
. Most protists are single-celled and microscopic, but there are exceptions, and some marine protists are neither single-celled nor microscopic, such as
seaweed Seaweed, or macroalgae, refers to thousands of species of macroscopic, multicellular, marine algae. The term includes some types of ''Rhodophyta'' (red), '' Phaeophyta'' (brown) and ''Chlorophyta'' (green) macroalgae. Seaweed species such as ...
.


Silicon-based shells

Although silicon is readily available in the form of
silicate A silicate is any member of a family of polyatomic anions consisting of silicon and oxygen, usually with the general formula , where . The family includes orthosilicate (), metasilicate (), and pyrosilicate (, ). The name is also used ...
s, very few organisms use it directly.
Diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s,
radiolaria The Radiolaria, also called Radiozoa, are unicellular eukaryotes of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ect ...
, and
siliceous sponge The siliceous sponges form a major clade of the phylum Porifera, consisting of classes Demospongiae (common sponges) and Hexactinellida (glass sponges). They are characterized by spicules made out of silicon dioxide, unlike calcareous sponges. ...
s use biogenic silica as a structural material for their skeletons. In more advanced plants, the silica
phytolith Phytoliths (from Greek language, Greek, "plant stone") are rigid, microscopic mineral deposits found in some plant tissues, often persisting after the decay of the plant. Although some use "phytolith" to refer to all mineral secretions by plants, ...
s (opal phytoliths) are rigid microscopic bodies occurring in the cell; some plants, including
rice Rice is a cereal grain and in its Domestication, domesticated form is the staple food of over half of the world's population, particularly in Asia and Africa. Rice is the seed of the grass species ''Oryza sativa'' (Asian rice)—or, much l ...
, need silica for their growth. Silica has been shown to improve plant cell wall strength and structural integrity in some plants.


Diatoms

Diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s form a (disputed) phylum containing about 100,000 recognised species of mainly unicellular algae. Diatoms generate about 20 per cent of the oxygen produced on the planet each year, take in over 6.7 billion metric tons of
silicon Silicon is a chemical element; it has symbol Si and atomic number 14. It is a hard, brittle crystalline solid with a blue-grey metallic lustre, and is a tetravalent metalloid (sometimes considered a non-metal) and semiconductor. It is a membe ...
each year from the waters in which they live, and contribute nearly half of the organic material found in the oceans. Diatoms are enclosed in protective silica (glass) shells called frustules. The beautifully engineered and intricate structure of many of these frustules is such that they are often referred to as "jewels of the sea". Each frustule is made from two interlocking parts covered with tiny holes through which the diatom exchanges nutrients and wastes.Wassilieff, Maggy (2006
"Plankton - Plant plankton"
''Te Ara - the Encyclopedia of New Zealand''. Accessed: 2 November 2019.
The frustules of dead diatoms drift to the ocean floor where, over millions of years, they can build up as much as half a mile deep. Diatoms uses silicon in the biogenic silica (BSiO2) form, which is taken up by the silicon
transport protein A transport protein (variously referred to as a transmembrane pump, transporter, escort protein, acid transport protein, cation transport protein, or anion transport protein) is a protein that serves the function of moving other materials within ...
to be predominantly used in constructing these protective cell wall structures. Silicon enters the ocean in a dissolved form such as
silicic acid In chemistry, a silicic acid () is any chemical compound containing the element silicon attached to oxide () and hydroxyl () groups, with the general formula or, equivalently, . Orthosilicic acid is a representative example. Silicic acids are ra ...
or
silicate A silicate is any member of a family of polyatomic anions consisting of silicon and oxygen, usually with the general formula , where . The family includes orthosilicate (), metasilicate (), and pyrosilicate (, ). The name is also used ...
. Since diatoms are one of the main users of these forms of silicon, they contribute greatly to the concentration of silicon throughout the ocean. Silicon forms a nutrient-like profile in the ocean due to the diatom productivity in shallow depths, which means there is less concentration of silicon in the upper ocean and more concentration of silicon in the deep ocean. Diatom productivity in the upper ocean contribute to the amount of silicon exported to the lower ocean. When diatom cells are lysed in the upper ocean, their nutrients like, iron, zinc, and silicon, are brought to the lower ocean through a process called
marine snow In the deep ocean, marine snow (also known as "ocean dandruff") is a continuous shower of mostly organic detritus falling from the upper layers of the water column. It is a significant means of exporting energy from the light-rich photic zone to ...
. Marine snow involves the downward transfer of
particulate organic matter Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 millimeters (53 μm) to 2 millimeters. Particulate org ...
by vertical mixing of
dissolved organic matter Dissolved organic carbon (DOC) is the fraction of organic carbon Operational definition, operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometre, micrometers. The fraction remain ...
. Availability of silicon appears crucial for diatom productivity, and as long as silicic acid is available for diatoms to utilize, the diatoms contribute other important nutrient concentrations in the deep ocean. In coastal zones, diatoms serve as the major
phytoplanktonic Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Phyt ...
organisms and greatly contribute to biogenic silica production. In the open ocean, however, diatoms have a reduced role in global annual silica production. Diatoms in North Atlantic and North Pacific subtropical gyres contribute only about 6% of global annual marine silica production, while the Southern Ocean produces about one-third of the global marine biogenic silica. The Southern Ocean is referred to as having a "biogeochemical divide", since only minuscule amounts of silicon is transported out of this region. File:Diatoms (248 05) Various diatoms.jpg,
Diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s are one of the most common types of phytoplankton File:Diatom Helipelta metil.jpg, Their protective shells (frustles) are made of silicon File:Diatom - Triceratium favus.jpg File:Diatom2.jpg, They come in many shapes and sizes
File:Diatom algae Amphora sp.jpg, Silicified frustule of a pennate diatom with two overlapping halves File:Fjouenne sbrmvr012w 20070924163039 small.jpg, ''Guinardia delicatula'', a diatom responsible for
algal bloom An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in fresh water or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term ''algae'' encompass ...
s in the North Sea and the English Channel File:Ископаемая диатомовая водоросль.jpg, Fossil diatom File:Pinnularia major.jpg, There are over 100,000 species of
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s which account for 50% of the ocean's primary production
Diatom frustules have been accumulating for over 100 million years, leaving rich deposits of nano and microstructured silicon oxide in the form of
diatomaceous earth Diatomaceous earth ( ), also known as diatomite ( ), celite, or kieselguhr, is a naturally occurring, soft, siliceous rock, siliceous sedimentary rock that can be crumbled into a fine white to off-white powder. It has a particle size ranging fr ...
around the globe. The evolutionary causes for the generation of nano and microstructured silica by photosynthetic algae are not yet clear. However, in 2018 it was shown that absorption of
ultraviolet light Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of th ...
by nanostructured silica protects the
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
in the algal cells, and this may be an evolutionary cause for the formation of the glass cages.De Tommasi, E., Congestri, R., Dardano, P., De Luca, A.C., Managò, S., Rea, I. and De Stefano, M. (2018) "UV-shielding and wavelength conversion by centric diatom nanopatterned frustules". ''Scientific Reports'', 8(1): 1–14. . Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License


Radiolarians

Radiolarian The Radiolaria, also called Radiozoa, are unicellular eukaryotes of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ecto ...
s are unicellular predatory
protists A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...
encased in elaborate globular shells (or "capsules"), 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 A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
, 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 crystals File:Spherical radiolarian 2.jpg, Cutaway schematic diagram of a spherical radiolarian shell File:Cladococcus abietinus.jpg, ''Cladococcus abietinus''


Calcium-based shells


Coccolithophores

Coccolithophore Coccolithophores, or coccolithophorids, are single-celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdom ...
s are minute unicellular photosynthetic protists with two flagella for locomotion. Most of them are protected by a shell called a
coccosphere Coccolithophores, or coccolithophorids, are single-celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdo ...
. Coccospheres are covered with ornate circular plates or scales called
coccolith Coccoliths are individual plates or scales of calcium carbonate formed by coccolithophores (single-celled phytoplankton such as ''Emiliania huxleyi'') and cover the cell surface arranged in the form of a spherical shell, called a '' coccosphere'' ...
s. The coccoliths are made from calcium carbonate. The term coccolithophore derives from the Greek for a ''seed carrying stone'', referring to their small size and the coccolith stones they carry. Under the right conditions they bloom, like other phytoplankton, and can turn the ocean milky white. File:Coccolithus pelagicus.jpg File:JRYSEM-247-05-azurapl.jpg,
Coccolithophores Coccolithophores, or coccolithophorids, are single-celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingd ...
named after the BBC documentary series '' The Blue Planet'' File:Emiliania huxleyi coccolithophore (PLoS).png, The
coccolithophore Coccolithophores, or coccolithophorids, are single-celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdom ...
''
Emiliania huxleyi ''Gephyrocapsa huxleyi'', also called ''Emiliania huxleyi'', is the most abundant species of coccolithophore in modern oceans found in almost all ecosystems from the equator to sub-polar regions, and from nutrient rich upwelling zones to nutr ...
''
There are benefits for protists that carry protective shells. The diagram on the left below shows some benefits coccolithophore get from carrying coccoliths. In the diagram, (A) represents accelerated photosynthesis including carbon concentrating mechanisms (CCM) and enhanced light uptake via scattering of scarce photons for deep-dwelling species. (B) represents protection from photodamage including sunshade protection from
ultraviolet light Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of th ...
(UV) and photosynthetic active radiation (PAR) and energy dissipation under high-light conditions. (C) represents armour protection includes protection against viral/bacterial infections and grazing by selective and nonselective grazers.Monteiro, F.M., Bach, L.T., Brownlee, C., Bown, P., Rickaby, R.E., Poulton, A.J., Tyrrell, T., Beaufort, L., Dutkiewicz, S., Gibbs, S. and Gutowska, M.A. (2016) "Why marine phytoplankton calcify". ''Science Advances'', 2(7): e1501822. . Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
There are also costs for protists that carry protective shells. The diagram on the right above shows some of the energetic costs coccolithophore incur from carrying coccoliths. In the diagram, the energetic costs are reported in percentage of total photosynthetic budget. (A) represents transport processes include the transport into the cell from the surrounding seawater of primary calcification substrates Ca2+ and HCO3− (black arrows) and the removal of the end product H+ from the cell (gray arrow). The transport of Ca2+ through the cytoplasm to the coccolith vesicle (CV) is the dominant cost associated with calcification. (B) represents metabolic processes include the synthesis of coccolith-associated
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long-chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wat ...
s (CAPs – gray rectangles) by the
Golgi complex The Golgi apparatus (), also known as the Golgi complex, Golgi body, or simply the Golgi, is an organelle found in most eukaryotic Cell (biology), cells. Part of the endomembrane system in the cytoplasm, it protein targeting, packages proteins ...
(white rectangles) that regulate the
nucleation In thermodynamics, nucleation is the first step in the formation of either a new Phase (matter), thermodynamic phase or Crystal structure, structure via self-assembly or self-organization within a substance or mixture. Nucleation is typically def ...
and
geometry Geometry (; ) is a branch of mathematics concerned with properties of space such as the distance, shape, size, and relative position of figures. Geometry is, along with arithmetic, one of the oldest branches of mathematics. A mathematician w ...
of CaCO3 crystals. The completed coccolith (gray plate) is a complex structure of intricately arranged CAPs and CaCO3 crystals. (C) Mechanical and structural processes account for the secretion of the completed coccoliths that are transported from their original position adjacent to the nucleus to the cell periphery, where they are transferred to the surface of the cell.


Foraminiferans

Like radiolarians,
foraminifera Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
ns (''forams'' for short) are single-celled predatory protists, also protected with shells that have holes in them. Their name comes from the Latin for "hole bearers". Their shells, often called
tests 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) ...
, may be single-chambered or multi-chambered; multi-chambered forams add more chambers as they grow. The most famous of these are made of calcite, but tests may also be made of
aragonite Aragonite is a carbonate mineral and one of the three most common naturally occurring crystal forms of calcium carbonate (), the others being calcite and vaterite. It is formed by biological and physical processes, including precipitation fr ...
, agglutinated sediment particles,
chiton Chitons () are marine molluscs of varying size in the class Polyplacophora ( ), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized. They are also sometimes known as sea cradles or coat-of-mail shells or suck ...
, or (rarely) of silica. Most forams are
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
, but about 40 living species are planktic. They are widely researched with well established fossil records which allow scientists to infer a lot about past environments and climates. Some foraminifera lack tests altogether. File:EB1911 Foraminifera - Section of Rotalia beccarii.jpg, section showing chambers of a spiral foram File:Live Ammonia tepida.jpg, Live '' Ammonia tepida'' streaming granular ectoplasm for catching food File:Planktic Foraminifera of the northern Gulf of Mexico.jpg, Group of planktonic forams File:Nummulitids.jpg, Fossil nummulitid forams of various sizes from the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
File:All Gizah Pyramids.jpg, The Egyptian pyramids were constructed from limestone that contained nummulites.


Other shells

The cell body of many
choanoflagellate Choanoflagellates are a group of free-living unicellular and colonial flagellate eukaryotes considered to be the closest living relatives of animals. The name refers to the characteristic funnel-shaped "collar" of interconnected microvilli and ...
s is surrounded by a distinguishing
extracellular matrix In biology, the extracellular matrix (ECM), also called intercellular matrix (ICM), is a network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide structural and bio ...
or
periplast The periplast is one of three types of cell-covering of three classes of algae. The cryptomonads have the periplast covering. The Dinophyceae have a type called the amphiesma, and the ''Euglena'' covering is the pellicle. Structure The periplast ...
. These cell coverings vary greatly in structure and composition and are used by taxonomists for classification purposes. Many choanoflagellates build complex basket-shaped "houses", called lorica, from several silica strips cemented together. The functional significance of the periplast is unknown, but in sessile organisms, it is thought to aid attachment to the substrate. In planktonic organisms, there is speculation that the periplast increases drag, thereby counteracting the force generated by the flagellum and increasing feeding efficiency. File:Salpingoeca sp..jpg,
Choanoflagellate Choanoflagellates are a group of free-living unicellular and colonial flagellate eukaryotes considered to be the closest living relatives of animals. The name refers to the characteristic funnel-shaped "collar" of interconnected microvilli and ...


Microfossils and sediments

The shells or skeletons of many protists survive over geological time scales as microfossils.
Microfossil A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
s are
fossils 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 ...
that are generally between 0.001mm and 1 mm in size, the study of which requires the use of light or electron
microscopy Microscopy is the technical field of using microscopes to view subjects too small to be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of microscopy: optical mic ...
. Fossils which can be studied by the naked eye or low-powered magnification, such as a hand lens, are referred to as macrofossils. Microfossils are a common feature of the geological record, from the
Precambrian The Precambrian ( ; or pre-Cambrian, sometimes abbreviated pC, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of t ...
to the
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
. They are most common in
marine sediment Marine sediment, or ocean sediment, or seafloor sediment, are deposits of insoluble particles that have accumulated on the seafloor. These particles either have their origins in soil and Rock (geology), rocks and have been Sediment transport, ...
s, but also occur in brackish water, fresh water and terrestrial
sedimentary Sedimentary rocks are types of rock formed by the cementation of sediments—i.e. particles made of minerals (geological detritus) or organic matter (biological detritus)—that have been accumulated or deposited at Earth's surface. Sedime ...
deposits. While every kingdom of
life Life, also known as biota, refers to matter that has biological processes, such as Cell signaling, signaling and self-sustaining processes. It is defined descriptively by the capacity for homeostasis, Structure#Biological, organisation, met ...
is represented in the microfossil record, the most abundant forms are
protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...
skeletons or
cysts A cyst is a closed Wikt:sac, sac, having a distinct Cell envelope, envelope and cell division, division compared with the nearby Biological tissue, tissue. Hence, it is a cluster of Cell (biology), cells that have grouped together to form a sac ...
from the
Chrysophyta Chrysophyta or golden algae is a term used to refer to certain heterokonts. It can be used to refer to: * Chrysophyceae (golden algae), Bacillariophyceae (diatoms), and Xanthophyceae (yellow-green algae) together. E.g., Adolf A. Pascher, Pascher ( ...
,
Pyrrhophyta The Dinoflagellates (), also called Dinophytes, are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered protists. Dinoflagellates are mostly marine plankton, but they are also commo ...
, Sarcodina,
acritarch Acritarchs () are organic microfossils, known from the Precambrian to the present. The classification is a catch all term used to refer to any organic microfossils that cannot be assigned to other groups. Their diversity reflects major ecolog ...
s and
chitinozoa Chitinozoa (singular: chitinozoan, plural: chitinozoans) are a group of flask-shaped, organic walled marine microfossils produced by an as yet unknown organism. Common from the Ordovician to Devonian periods (i.e. the mid-Paleozoic), the millim ...
ns, together with
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
and
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 ...
from the
vascular plant Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
s. In 2017, fossilized
microorganism A microorganism, or microbe, is an organism of microscopic scale, microscopic size, which may exist in its unicellular organism, single-celled form or as a Colony (biology)#Microbial colonies, colony of cells. The possible existence of unseen ...
s, or microfossils, were discovered in
hydrothermal vent Hydrothermal vents are fissures on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hot ...
precipitates in the
Nuvvuagittuq Belt The Nuvvuagittuq Greenstone Belt (NGB; Inuktitut: ) is a sequence of metamorphosed mafic to ultramafic volcanic and associated sedimentary rocks (a greenstone belt) located on the eastern shore of Hudson Bay, 40 km southeast of Inukjuak, Qu ...
that may be as old as 4.28 billion years old, the oldest record of life on Earth, suggesting "an almost instantaneous emergence of life" (in a geological time-scale sense), after ocean formation 4.41 billion years ago, and not long after the formation of the Earth 4.54 billion years ago. Nonetheless, life may have started even earlier, at nearly 4.5 billion years ago, as claimed by some researchers.


See also

*
Cytoskeleton The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compos ...
* Particulate inorganic matter


References


Further references

* Xu, K., Hutchins, D. and Gao, K. (2018) "Coccolith arrangement follows Eulerian mathematics in the coccolithophore ''Emiliania huxleyi''". ''PeerJ'', 6: e4608. {{doi, 10.1126/science.aaa7378.
Protistan Skeletons: A Geologic History of Evolution and Constraint
Protists Biomineralization Skeletal system