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Many protists have protective shells or tests, usually made from (glass) or calcium carbonate (chalk). are a diverse group of organisms that are not plants, animals, or fungi. They are typically microscopic unicellular organisms 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 . Although protists are typically very small, they are ubiquitous. Their numbers are such that their shells play a huge part in the formation of and in the global cycling of elements and nutrients.

The role of protist shells depends on the type of protist. Protists such as and have intricate, glass-like shells made of 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. and also have hard protective shells, but the shells are made of calcium carbonate. These shells can help with buoyancy, allowing the organisms to float in the 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 (including modern bacteria and archaea) and later evolved into . Eukaryotes include organisms such as plants, animals, fungi and "protists". Protists are usually single-celled and microscopic. They can be , meaning they obtain nutrients by consuming other organisms, or , meaning they produce their own food through or , or , 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 group that does not contain all descendants of a common ancestor. As such it does not constitute a and is not currently in formal scientific use. Nonetheless, the term continues to be used informally to refer to those 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 , as well as such as and . Many protists are asexual but can reproduce rapidly through or by fragmentation; others (including ) may reproduce either sexually or asexually.

(1982). 9780910249058, University of Tennessee, Dept. of Geological Sciences.

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 . Most protists are single-celled and microscopic, but there are exceptions, and some are neither single-celled nor microscopic, such as .


Silicon-based shells
Although silicon is readily available in the form of , very few organisms use it directly. , , and use as a structural material for their skeletons. In more advanced plants, the silica (opal phytoliths) are rigid microscopic bodies occurring in the cell; some plants, including , need silica for their growth.
(2026). 9780444531810, Elsevier Science.
Silica has been shown to improve plant cell wall strength and structural integrity in some plants.


Diatoms
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, The Air You're Breathing? A Diatom Made That take in over 6.7 billion metric tons of 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 . The beautifully engineered and intricate structure of many of these frustules is such that they are often referred to as "jewels of the sea".Ireland, T., "Engineering with algae". Biologist, 63(5): 10. 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 (BSiO2) form, which is taken up by the silicon transport protein to be predominantly used in constructing these protective cell wall structures. Silicon enters the ocean in a dissolved form such as or . 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 in the upper ocean, their nutrients like, iron, zinc, and silicon, are brought to the lower ocean through a process called . Marine snow involves the downward transfer of particulate organic matter by vertical mixing of dissolved organic matter. 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 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| 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 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 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 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 by nanostructured silica protects the 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 a Creative Commons Attribution 4.0 International License.


Radiolarians
are unicellular predatory protists 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, 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 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
are minute unicellular photosynthetic protists with two flagella for locomotion. Most of them are protected by a shell called a . Coccospheres are covered with ornate circular plates or scales called . 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.Wassilieff, Maggy (2006) "A coccolithophore", Te Ara - the Encyclopedia of New Zealand. Accessed: 2 November 2019.

File:Coccolithus pelagicus.jpg File:JRYSEM-247-05-azurapl.jpg| Coccolithophores named after the BBC documentary series The Blue Planet File:Emiliania huxleyi coccolithophore (PLoS).png|The Emiliania huxleyi

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 (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 a 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 (CAPs – gray rectangles) by the (white rectangles) that regulate the and geometry 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, ( 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, 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 , agglutinated sediment particles, , or (rarely) of silica. Most forams are , but about 40 living species are .
(1989). 9783540968153, Springer-Verlag. .
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 streaming granular ectoplasm for catching food File:Planktic Foraminifera of the northern Gulf of Mexico.jpg|Group of planktonic forams File:Nummulitids.jpg|Fossil forams of various sizes from the File:All Gizah Pyramids.jpg|The were constructed from limestone that contained . Foraminifera: History of Study, University College London. Retrieved: 18 November 2019.


Other shells
The cell body of many is surrounded by a distinguishing extracellular matrix or . 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.

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Microfossils and sediments
The shells or skeletons of many protists survive over geological time scales as microfossils. are that are generally between 0.001mm and 1 mm in size, the study of which requires the use of light or electron . Fossils which can be studied by the naked eye or low-powered magnification, such as a hand lens, are referred to as .

Microfossils are a common feature of the geological record, from the to the . They are most common in , but also occur in brackish water, fresh water and terrestrial deposits. While every kingdom of is represented in the microfossil record, the most abundant forms are skeletons or from the , , , and , together with and from the .

In 2017, fossilized , or microfossils, were discovered in hydrothermal vent in the Nuvvuagittuq Belt 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


Further references

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