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 .
Coccolithophores are spherical cells about 5–100 micrometres across, enclosed by calcareous plates called coccoliths, which are about 2–25 micrometres across. [1] Coccolithophores are an important group of about 200 marine phytoplankton species [2] which cover themselves with a calcium carbonate shell called a "coccosphere". They are ecologically and biogeochemically important but the reason why they calcify remains elusive. One key function may be that the coccosphere offers protection against microzooplankton predation, which is one of the main causes of phytoplankton death in the ocean. [3]
Coccolithophores have been an integral part of marine plankton communities since the Jurassic. [7] [8] Today, coccolithophores contribute ~1–10% to primary production in the surface ocean [9] and ~50% to pelagic CaCO3 sediments. [10] Their calcareous shell increases the sinking velocity of photosynthetically fixed CO2 into the deep ocean by ballasting organic matter. [11] [12] At the same time, the biogenic precipitation of calcium carbonate during coccolith formation reduces the total alkalinity of seawater and releases CO2. [13] [14] Thus, coccolithophores play an important role in the marine carbon cycle by influencing the efficiency of the biological carbon pump and the oceanic uptake of atmospheric CO2. [3]
As of 2021, it is not known why coccolithophores calcify and how their ability to produce coccoliths is associated with their ecological success. [15] [16] [17] [18] [19] The most plausible benefit of having a coccosphere seems to be a protection against predators or viruses. [20] [18] Viral infection is an important cause of phytoplankton death in the oceans, [21] and it has recently been shown that calcification can influence the interaction between a coccolithophore and its virus. [22] [23] The major predators of marine phytoplankton are microzooplankton like ciliates and dinoflagellates. These are estimated to consume about two-thirds of the primary production in the ocean [24] and microzooplankton can exert a strong grazing pressure on coccolithophore populations. [25] Although calcification does not prevent predation, it has been argued that the coccosphere reduces the grazing efficiency by making it more difficult for the predator to utilise the organic content of coccolithophores. [26] Heterotrophic protists are able to selectively choose prey on the basis of its size or shape and through chemical signals [27] [28] and may thus favor other prey that is available and not protected by coccoliths. [3]
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Coccoliths are formed within the cell in vesicles derived from the golgi body. When the coccolith is complete these vesicles fuse with the cell wall and the coccolith is exocytosed and incorporated in the coccosphere. The coccoliths are either dispersed following death and breakup of the coccosphere, or are shed continually by some species. They sink through the water column to form an important part of the deep-sea sediments (depending on the water depth). Thomas Huxley was the first person to observe these forms in modern marine sediments and he gave them the name 'coccoliths' in a report published in 1858. [29] [30] Coccoliths are composed of calcium carbonate as the mineral calcite and are the main constituent of chalk deposits such as the white cliffs of Dover (deposited in Cretaceous times), in which they were first described by Henry Clifton Sorby in 1861. [31]
There are two main types of coccoliths, heterococcoliths and holococcoliths. Heterococcoliths are formed of a radial array of elaborately shaped crystal units. Holococcoliths are formed of minute (~0.1 micrometre) calcite rhombohedra, arranged in continuous arrays. The two coccolith types were originally thought to be produced by different families of coccolithophores. Now, however, it is known through a mix of observations on field samples and laboratory cultures, that the two coccolith types are produced by the same species but at different life cycle phases. Heterococcoliths are produced in the diploid life-cycle phase and holococcoliths in the haploid phase. Both in field samples and laboratory cultures, there is the possibility of observing a cell covered by a combination of heterococcoliths and holococcoliths. This indicates the transition from the diploid to the haploid phase of the species. Such combination of coccoliths has been observed in field samples, with many of them coming from the Mediterranean. [32] [33]
Coccoliths are also classified depending on shape. Common shapes include: [34] [35]
Although coccoliths are remarkably elaborate structures whose formation is a complex product of cellular processes, their function is unclear. Hypotheses include defence against grazing by zooplankton or infection by bacteria or viruses; maintenance of buoyancy; release of carbon dioxide for photosynthesis; to filter out harmful UV light; or in deep-dwelling species, to concentrate light for photosynthesis.
Because coccoliths are formed of low-Mg calcite, the most stable form of calcium carbonate, they are readily fossilised. They are found in sediments together with similar microfossils of uncertain affinities (nanoliths) from the Upper Triassic to recent. They are widely used as biostratigraphic markers and as paleoclimatic proxies. Stable oxygen and carbon isotope data from coccoliths is used to reconstruct estimates of oceanic CO2 concentrations in the geologic past. [37] Coccoliths and related fossils are referred to as calcareous nanofossils or calcareous nannoplankton (nanoplankton) .
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 Protista, according to Robert Whittaker's five-kingdom system, or clade Hacrobia, according to a newer biological classification system. Within the Hacrobia, the coccolithophores are in the phylum or division Haptophyta, class Prymnesiophyceae. Coccolithophores are almost exclusively marine, are photosynthetic and mixotrophic, and exist in large numbers throughout the sunlight zone of the ocean.
The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sediments. In other words, it is a biologically mediated process which results in the sequestering of carbon in the deep ocean away from the atmosphere and the land. The biological pump is the biological component of the "marine carbon pump" which contains both a physical and biological component. It is the part of the broader oceanic carbon cycle responsible for the cycling of organic matter formed mainly by phytoplankton during photosynthesis (soft-tissue pump), as well as the cycling of calcium carbonate (CaCO3) formed into shells by certain organisms such as plankton and mollusks (carbonate pump).
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, such as a hand lens, is referred to as a macrofossil.
Gephyrocapsa huxleyi, formerly called Emiliania huxleyi, is a species of coccolithophore found in almost all ocean ecosystems from the equator to sub-polar regions, and from nutrient rich upwelling zones to nutrient poor oligotrophic waters. It is one of thousands of different photosynthetic plankton that freely drift in the photic zone of the ocean, forming the basis of virtually all marine food webs. It is studied for the extensive blooms it forms in nutrient-depleted waters after the reformation of the summer thermocline. Like other coccolithophores, E. huxleyi is a single-celled phytoplankton covered with uniquely ornamented calcite disks called coccoliths. Individual coccoliths are abundant in marine sediments although complete coccospheres are more unusual. In the case of E. huxleyi, not only the shell, but also the soft part of the organism may be recorded in sediments. It produces a group of chemical compounds that are very resistant to decomposition. These chemical compounds, known as alkenones, can be found in marine sediments long after other soft parts of the organisms have decomposed. Alkenones are most commonly used by earth scientists as a means to estimate past sea surface temperatures.
Ocean acidification is the ongoing decrease in the pH of the Earth's ocean. Between 1950 and 2020, the average pH of the ocean surface fell from approximately 8.15 to 8.05. Carbon dioxide emissions from human activities are the primary cause of ocean acidification, with atmospheric carbon dioxide levels exceeding 410 ppm. CO2 from the atmosphere is absorbed by the oceans. This chemical reaction produces carbonic acid which dissociates into a bicarbonate ion and a hydrogen ion. The presence of free hydrogen ions lowers the pH of the ocean, increasing acidity. Marine calcifying organisms, such as mollusks and corals, are especially vulnerable because they rely on calcium carbonate to build shells and skeletons.
Calcareous is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines.
The carbonate compensation depth (CCD) is the depth, in the oceans, at which the rate of supply of calcium carbonates matches the rate of solvation. That is, solvation 'compensates' supply. Below the CCD solvation is faster, so that carbonate particles dissolve and the carbonate shells (tests) of animals are not preserved. Carbonate particles cannot accumulate in the sediments where the sea floor is below this depth.
Alkenones are long-chain unsaturated methyl and ethyl n-ketones produced by a few phytoplankton species of the class Prymnesiophyceae. Alkenones typically contain between 35 and 41 carbon atoms and with between two and four double bonds. Uniquely for biolipids, alkenones have a spacing of five methylene groups between double bonds, which are of the less common E configuration. The biological function of alkenones remains under debate although it is likely that they are storage lipids. Alkenones were first described in ocean sediments recovered from Walvis Ridge and then shortly afterwards in cultures of the marine coccolithophore Gephyrocapsa huxleyi. The earliest known occurrence of alkenones is during the Aptian 120 million years ago. They are used in organic geochemistry as a proxy for past sea surface temperature.
Pleurochrysis carterae is a marine species of unicellular coccolithophorid algae that has the ability to calcify subcellularly. They produce calcified scales, known as coccoliths, which are deposited on the surface of the cell resulting in the formation of a coccosphere. Pleurochrysis carterae produce heterococcoliths which are composed of crystal units of variable shapes and sizes.
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 sea, mainly by rivers but also by dust carried by wind and by the flow of glaciers into the sea, or they are biogenic deposits from marine organisms or from chemical precipitation in seawater, as well as from underwater volcanoes and meteorite debris.
Calanus finmarchicus is a species of copepod and a component of the zooplankton, which is found in enormous amounts in the northern Atlantic Ocean.
Emiliania is a global coccolithophorid genus.
The oceanic carbon cycle is composed of processes that exchange carbon between various pools within the ocean as well as between the atmosphere, Earth interior, and the seafloor. The carbon cycle is a result of many interacting forces across multiple time and space scales that circulates carbon around the planet, ensuring that carbon is available globally. The Oceanic carbon cycle is a central process to the global carbon cycle and contains both inorganic carbon and organic carbon. Part of the marine carbon cycle transforms carbon between non-living and living matter.
Marine biogenic calcification is the production of calcium carbonate by organisms in the global ocean.
Rappephyceae, or Rappemonads, are a small family of protists first described in 2011, of uncertain phylogenic affinity. It has been discussed as a possible member of a larger clade Haptophyta. This newly identified taxonomic class of phytoplankton are named after a professor from the Hawai’i institute of marine biology, known as Michael Rappé. Rappé discovered these phytoplankton in the Atlantic Ocean and published his findings on their DNA in 1998. Current research has shown that these organisms provide an immense amount of nutritional molecules, such as oxygen, for other organisms using biochemical processes like photosynthesis and carbon fixation.
Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as marine single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics because they are paraphyletic.
Many protists have protective shells or tests, usually made from silica (glass) or calcium carbonate (chalk). Protists are a diverse group of eukaryote organisms that are not plants, animals, or fungi. They are typically microscopic unicellular organisms that live in water or moist environments.
Particulate inorganic carbon (PIC) can be contrasted with dissolved inorganic carbon (DIC), the other form of inorganic carbon found in the ocean. These distinctions are important in chemical oceanography. Particulate inorganic carbon is sometimes called suspended inorganic carbon. In operational terms, it is defined as the inorganic carbon in particulate form that is too large to pass through the filter used to separate dissolved inorganic carbon.
The Great Calcite Belt (GCB) refers to a region of the ocean where there are high concentrations of calcite, a mineral form of calcium carbonate. The belt extends over a large area of the Southern Ocean surrounding Antarctica. The calcite in the Great Calcite Belt is formed by tiny marine organisms called coccolithophores, which build their shells out of calcium carbonate. When these organisms die, their shells sink to the bottom of the ocean, and over time, they accumulate to form a thick layer of calcite sediment.
Calcareous nannofossils are a class of tiny microfossils that are similar to coccoliths deposited by the modern-day coccolithophores. The nannofossils are a convenient source of geochronological data due to the abundance and rapid evolution of the single-cell organisms forming them (nannoplankton) and ease of handling of the sediment samples. The practical applications of calcareous nannofossils in the areas of biostratigraphy and paleoecology became clear once the deepwater drilling took off in 1968 with the Deep Sea Drilling Project, and they have been extensively studied ever since. Nannofossils provide one of the most important paleontological records with the contiguous length of 220 million years.
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