Coccolithophore - Knowledge

2167:. During photosynthesis carbon dioxide is removed from the water, making it more basic. Also calcification removes carbon dioxide, but chemistry behind it leads to the opposite pH reaction; it makes the water more acidic. The combination of photosynthesis and calcification therefore even out each other regarding pH changes. In addition, these exoskeletons may confer an advantage in energy production, as coccolithogenesis seems highly coupled with photosynthesis. Organic precipitation of calcium carbonate from bicarbonate solution produces free carbon dioxide directly within the cellular body of the alga, this additional source of gas is then available to the Coccolithophore for photosynthesis. It has been suggested that they may provide a cell-wall like barrier to isolate intracellular chemistry from the marine environment. More specific, defensive properties of coccoliths may include protection from osmotic changes, chemical or mechanical shock, and short-wavelength light. It has also been proposed that the added weight of multiple layers of coccoliths allows the organism to sink to lower, more nutrient rich layers of the water and conversely, that coccoliths add buoyancy, stopping the cell from sinking to dangerous depths. Coccolith appendages have also been proposed to serve several functions, such as inhibiting grazing by zooplankton. 601: 2400:) plates that constitute the dinoflagellate shell, should rather be favored at high H concentrations because these usually coincide with high . Under these conditions dinoflagellates could down-regulate the energy-consuming operation of carbon concentrating mechanisms to fuel the production of organic source material for their shell. Therefore, a shift in carbonate chemistry conditions toward high may promote their competitiveness relative to coccolithophores. However, such a hypothetical gain in competitiveness due to altered carbonate chemistry conditions would not automatically lead to dinoflagellate dominance because a huge number of factors other than carbonate chemistry have an influence on 2196: 2295: 1322: 700: 2497: 852: 1338: 7927: 3184: 143: 1036: 1446: 2980: 2028: 774:

coccolithophores are K strategist and are usually found on nutrient-poor surface waters. They are poor competitors when compared to other phytoplankton and thrive in habitats where other phytoplankton would not survive. These two stages in the life cycle of coccolithophores occur seasonally, where more nutrition is available in warmer seasons and less is available in cooler seasons. This type of life cycle is known as a complex heteromorphic life cycle.

783: 120: 951: 231: 6669: 6363: 5490: 5439: 5323: 3499: 3396: 585:. These are estimated to consume about two-thirds of the primary production in the ocean and microzooplankton can exert a strong grazing pressure on coccolithophore populations. 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. 3137:, impairs ion channel function and therefore places evolutionary selective pressure on coccolithophores and makes them (and other ocean calcifiers) vulnerable to ocean acidification. In 2008, field evidence indicating an increase in calcification of newly formed ocean sediments containing coccolithophores bolstered the first ever experimental data showing that an increase in ocean CO 2392:(diatom shell) seems to be the most inexpensive armor under all circumstances because diatoms typically outcompete all other groups when silicate is available. The coccosphere is relatively inexpensive under sufficient , high , and low because the substrate is saturating and protons are easily released into seawater. In contrast, the construction of 2282:(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. The costs associated with these processes are likely to be comparable to organic-scale 920: 890: 904: 808:

considered the Central North Zone which is an area between 30 N and 5 N, composed of the North Equatorial Current and the Equatorial Countercurrent. These two currents move in opposite directions, east and west, allowing for a strong mixing of waters and allowing a large variety of species to populate the area.

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have been found to produce haemolytic compounds, the agent responsible for toxicity. Some of these toxic species are responsible for large fish kills and can be accumulated in organisms such as shellfish; transferring it through the food chain. In laboratory tests for toxicity members of the oceanic

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Recent studies show that climate change has direct and indirect impacts on Coccolithophore distribution and productivity. They will inevitably be affected by the increasing temperatures and thermal stratification of the top layer of the ocean, since these are prime controls on their ecology, although

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is instead released back into the atmosphere. As a result of this, researchers have postulated that large blooms of coccolithophores may contribute to global warming in the short term. A more widely accepted idea, however, is that over the long term coccolithophores contribute to an overall decrease

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prey and found no evidence that the coccosphere prevents ingestion by the grazer. Instead, ingestion rates were dependent on the offered genotype of E. huxleyi. Altogether, these two studies suggest that the genotype has a strong influence on ingestion by the microzooplankton species, but if and how

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crystals. These crystals are thought to form at least partially outside the cell. Heterococcoliths occur only in the diploid phase, have radial symmetry, and are composed of relatively few complex crystal units (fewer than 100). Although they are rare, combination coccospheres, which contain both

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or other groups of phytoplankton, such as coccolithophores. A low silicate to nitrogen and phosphorus ratio allows coccolithophores to outcompete other phytoplankton species; however, when silicate to phosphorus to nitrogen ratios are high coccolithophores are outcompeted by diatoms. The increase in

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The upper photic zone is low in nutrient concentration, high in light intensity and penetration, and usually higher in temperature. The lower photic zone is high in nutrient concentration, low in light intensity and penetration and relatively cool. The middle photic zone is an area that contains the

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conditions, the most abundant areas of coccolithophores where there is the highest species diversity are located in subtropical zones with a temperate climate. While water temperature and the amount of light intensity entering the water's surface are the more influential factors in determining where

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Daniels, Chris J.; Poulton, Alex J.; Balch, William M.; Marañón, Emilio; Adey, Tim; Bowler, Bruce C.; Cermeño, Pedro; Charalampopoulou, Anastasia; Crawford, David W.; Drapeau, Dave; Feng, Yuanyuan; Fernández, Ana; Fernández, Emilio; Fragoso, Glaucia M.; González, Natalia; Graziano, Lisa M.; Heslop,

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saturation contrary to predictions. Understanding the effects of increasing ocean acidification on coccolithophore species is absolutely essential to predicting the future chemical composition of the ocean, particularly its carbonate chemistry. Viable conservation and management measures will come

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The diagram on the left shows the benefits of coccolithophore calcification. (A) Accelerated photosynthesis includes CCM (1) and enhanced light uptake via scattering of scarce photons for deep-dwelling species (2). (B) Protection from photodamage includes sunshade protection from ultraviolet (UV)

1349: (E), the reference species for coccolithophore studies, is contrasted with a range of other species spanning the biodiversity of modern coccolithophores. All images are scanning electron micrographs of cells collected by seawater filtration from the open ocean. Species illustrated: (A) 807:

Within the Pacific Ocean, approximately 90 species have been identified with six separate zones relating to different Pacific currents that contain unique groupings of different species of coccolithophores. The highest diversity of coccolithophores in the Pacific Ocean was in an area of the ocean

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and added to the inner surface of the coccosphere. This means that the most recently produced coccoliths may lie beneath older coccoliths. Depending upon the phytoplankton's stage in the life cycle, two different types of coccoliths may be formed. Holococcoliths are produced only in the haploid

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need sunlight and nutrients from the ocean to survive, so they thrive in areas with large inputs of nutrient rich water upwelling from the lower levels of the ocean. Most coccolithophores require sunlight only for energy production, and have a higher ratio of nitrate uptake over ammonium uptake

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The complete distribution of coccolithophores is currently not known and some regions, such as the Indian Ocean, are not as well studied as other locations in the Pacific and Atlantic Oceans. It is also very hard to explain distributions due to multiple constantly changing factors involving the

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of coccolithophores depend on their life cycle stage. When coccolithophores are diploid, they are r-selected. In this phase they tolerate a wider range of nutrient compositions. When they are haploid they are K- selected and are often more competitive in stable low nutrient environments. Most

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chemistry conditions is possible within one year. Unraveling these fundamental constraints and the limits of adaptation should be a focus in future coccolithophore studies because knowing them is the key information required to understand to what extent the calcification response to carbonate

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process known as coccolithogenesis. Generally, calcification of coccoliths occurs in the presence of light, and these scales are produced much more during the exponential phase of growth than the stationary phase. Although not yet entirely understood, the biomineralization process is tightly

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concentrations. During calcification two carbon atoms are taken up and one of them becomes trapped as calcium carbonate. This calcium carbonate sinks to the bottom of the ocean in the form of coccoliths and becomes part of sediment; thus, coccolithophores provide a sink for emitted carbon,

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As of 2021, it is not known why coccolithophores calcify and how their ability to produce coccoliths is associated with their ecological success. The most plausible benefit of having a coccosphere seems to be a protection against predators or viruses. Viral infection is an important cause of

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Balch, W. M.; Drapeau, D. T.; Bowler, B. C.; Lyczskowski, E.; Booth, E. S.; Alley, D. (2011). "The contribution of coccolithophores to the optical and inorganic carbon budgets during the Southern Ocean Gas Exchange Experiment: New evidence in support of the "Great Calcite Belt" hypothesis".

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help to produce thicker clouds to block the sun. When the oceans cool, the number of coccolithophorids decrease and the amount of clouds also decrease. When there are fewer clouds blocking the sun, the temperature also rises. This, therefore, maintains the balance and equilibrium of nature.

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predominance. The overlap of two major phytoplankton groups, coccolithophores and diatoms, in the dynamic frontal systems characteristic of this region provides an ideal setting to study environmental influences on the distribution of different species within these taxonomic groups.

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In this process the coccoliths from the parent cell are divided between the two daughter cells. There have been suggestions stating the possible presence of a sexual reproduction process due to the diploid stages of the coccolithophores, but this process has never been observed.

956: 1249:. These viruses, known as E. huxleyi viruses (EhVs), appear to infect the coccosphere coated diploid phase of the life cycle almost exclusively. It has been proposed that as the haploid organism is not infected and therefore not affected by the virus, the co-evolutionary " 2335:) to co-adapt in order to keep H efflux alive. The obligatory H efflux associated with calcification may therefore pose a fundamental constraint on adaptation which may potentially explain why "calcification crisis" were possible during long-lasting (thousands of years) CO 4328:

Monteiro, Fanny M.; Bach, Lennart T.; Brownlee, Colin; Bown, Paul; Rickaby, Rosalind E. M.; Poulton, Alex J.; Tyrrell, Toby; Beaufort, Luc; Dutkiewicz, Stephanie; Gibbs, Samantha; Gutowska, Magdalena A.; Lee, Renee; Riebesell, Ulf; Young, Jeremy; Ridgwell, Andy (2016).

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Although motility and colony formation vary according to the life cycle of different coccolithophore species, there is often alternation between a motile, haploid phase, and a non-motile diploid phase. In both phases, the organism's dispersal is largely due to ocean

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Rachel; Holligan, Patrick M.; Hopkins, Jason; Huete-Ortega, María; Hutchins, David A.; Lam, Phoebe J.; Lipsen, Michael S.; López-Sandoval, Daffne C.; Loucaides, Socratis; Marchetti, Adrian; Mayers, Kyle M. J.; Rees, Andrew P.; Sobrino, Cristina; et al. (2018).

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Their predators include the common predators of all phytoplankton including small fish, zooplankton, and shellfish larvae. Viruses specific to this species have been isolated from several locations worldwide and appear to play a major role in spring bloom dynamics.

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do not need to sustain the calcification-related H efflux. Thus, they probably do not need to adapt in order to keep costs for the production of structural elements low. On the contrary, dinoflagellates (except for calcifying species; with generally inefficient

1140:, meaning photosythetic production is diminished due to an excess of light. In case 1), a high concentration of coccoliths leads to a simultaneous increase in surface water temperature and decrease in the temperature of deeper waters. This results in more 790:

Coccolithophores occur throughout the world's oceans. Their distribution varies vertically by stratified layers in the ocean and geographically by different temporal zones. While most modern coccolithophores can be located in their associated stratified

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Mayers, Kyle M. J.; Poulton, Alex J.; Bidle, Kay; Thamatrakoln, Kimberlee; Schieler, Brittany; Giering, Sarah L. C.; Wells, Seona R.; Tarran, Glen A.; Mayor, Dan; Johnson, Matthew; Riebesell, Ulf; Larsen, Aud; Vardi, Assaf; Harvey, Elizabeth L. (2020).

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that cover up to 35% of the ocean floor and is kilometres thick in places. Because of their abundance and wide geographic ranges, the coccoliths which make up the layers of this ooze and the chalky sediment formed as it is compacted serve as valuable

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light and photosynthetic active radiation (PAR) (1) and energy dissipation under high-light conditions (2). (C) Armor protection includes protection against viral/bacterial infections (1) and grazing by selective (2) and nonselective (3) grazers.

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Honjo, Susumu; Manganini, Steven J.; Krishfield, Richard A.; Francois, Roger (2008). "Particulate organic carbon fluxes to the ocean interior and factors controlling the biological pump: A synthesis of global sediment trap programs since 1983".

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to produce haploid cells again, starting the cycle over. With coccolithophores, asexual reproduction by mitosis is possible in both phases of the life cycle, which is a contrast with most other organisms that have alternating life cycles. Both

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holococcoliths and heterococcoliths, have been observed in the plankton recording coccolithophore life cycle transitions. Finally, the coccospheres of some species are highly modified with various appendages made of specialized coccoliths.

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spring and summer in the Southern Ocean, plays an important role in climate fluctuations, accounting for over 60% of the Southern Ocean area (30–60° S). The region between 30° and 50° S has the highest uptake of anthropogenic carbon dioxide

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Currently, the evidence supporting or refuting a protective function of the coccosphere against predation is limited. Some researchers found that overall microzooplankton predation rates were reduced during blooms of the coccolithophore

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While the exact function of the coccosphere is unclear, many potential functions have been proposed. Most obviously coccoliths may protect the phytoplankton from predators. It also appears that it helps them to create a more stable

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Houdan; Probert, I; Zatylny, C; Véron, B; Billard, C; et al. (2006), ". Ecology of oceanic coccolithophores. I. Nutritional preferences of the two stages in the life cycle of Coccolithus braarudii and Calcidiscus leptoporus",

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Olson, M.Brady; Strom, Suzanne L. (2002). "Phytoplankton growth, microzooplankton herbivory and community structure in the southeast Bering Sea: Insight into the formation and temporal persistence of an Emiliania huxleyi bloom".

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Moheimani, N.R.; Webb, J.P.; Borowitzka, M.A. (2012), "Bioremediation and other potential applications of coccolithophorid algae: A review. . Bioremediation and other potential applications of coccolithophorid algae: A review",

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Lohbeck, Annette; Tietjens, Maike; Bund, Andreas (2014). "Das physische Selbstkonzept, die individuell präferierte Bezugsnormorientierung und die Zielorientierung bei Grundschulkindern der zweiten und vierten Jahrgangsstufe".

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Fu, Fei-Xue; Zhang, Yaohong; Warner, Mark E.; Feng, Yuanyuan; Sun, Jun; Hutchins, David A. (2008). "A comparison of future increased CO2 and temperature effects on sympatric Heterosigma akashiwo and Prorocentrum minimum".

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Poulton, Alex J.; Adey, Tim R.; Balch, William M.; Holligan, Patrick M. (2007). "Relating coccolithophore calcification rates to phytoplankton community dynamics: Regional differences and implications for carbon export".

2099:. The coccoliths are created inside the coccolithophore cell and while some species maintain a single layer throughout life only producing new coccoliths as the cell grows, others continually produce and shed coccoliths. 2318:

for intra-cellular calcification) to become more costly with ongoing ocean acidification as the electrochemical H inside-out gradient is reduced and passive proton outflow impeded. Adapted cells would have to activate

3129:, as coccolith production would otherwise produce a toxic excess of H ions. When the function of these ion channels is disrupted, the coccolithophores stop the calcification process to avoid acidosis, thus forming a 1148:

of the ocean is less than that from anthropogenic factors. Therefore, the overall result of large blooms of coccolithophores is a decrease in water column productivity, rather than a contribution to global warming.

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Boeckel; Baumann, Karl-Heinz; Henrich, Rüdiger; Kinkel, Hanno; et al. (2006), "Coccolith distribution patterns in South Atlantic and Southern Ocean surface sediments in relation to environmental gradients",

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Durak, G.M., Taylor, A.R., Walker, C.E., Probert, I., De Vargas, C., Audic, S., Schroeder, D., Brownlee, C. and Wheeler, G.L. (2016) "A role for diatom-like silicon transporters in calcifying coccolithophores".

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Sabine, C. L.; Feely, R. A.; Gruber, N.; Key, R. M.; Lee, K.; Bullister, J. L.; Wanninkhof, R.; Wong, C. S.; Wallace, D. W.; Tilbrook, B.; Millero, F. J.; Peng, T. H.; Kozyr, A.; Ono, T.; Rios, A. F. (2004).

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Hay, W.W.; Mohler, H.P.; Roth, P.H.; Schmidt, R.R.; Boudreaux, J.E. (1967), "Calcareous nannoplankton zonation of the Cenozoic of the Gulf Coast and Caribbean-Antillean area, and transoceanic correlation",

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fertilization. Under the assumption that any form of shell/exoskeleton protects phytoplankton against predation non-calcareous armors may be the preferable solution to realize protection in a future ocean.

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No environmental evidence of coccolithophore toxicity has been reported, but they belong to the class Prymnesiophyceae which contain orders with toxic species. Toxic species have been found in the genera

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phytoplankton death in the oceans, and it has recently been shown that calcification can influence the interaction between a coccolithophore and its virus. The major predators of marine phytoplankton are

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in the atmosphere may affect the calcification machinery of coccolithophores. This may not only affect immediate events such as increases in population or coccolith production, but also may induce

955: 1225:(nitrogen is required for growth and can be used directly from nitrate but not ammonium). Because of this they thrive in still, nutrient-poor environments where other phytoplankton are starving. 537:. Today, coccolithophores contribute ~1–10% to inorganic carbon fixation (calcification) to total carbon fixation (calcification plus photosynthesis) in the surface ocean and ~50% to pelagic CaCO 5456:

Smith, Helen E. K.; Poulton, Alex J.; Garley, Rebecca; Hopkins, Jason; Lubelczyk, Laura C.; Drapeau, Dave T.; Rauschenberg, Sara; Twining, Ben S.; Bates, Nicholas R.; Balch, William M. (2017).

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Fileman, E.S.; Cummings, D.G.; Llewellyn, C.A. (2002). "Microplankton community structure and the impact of microzooplankton grazing during an Emiliania huxleyi bloom, off the Devon coast".

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Sarmiento, J. L.; Slater, R.; Barber, R.; Bopp, L.; Doney, S. C.; Hirst, A. C.; Kleypas, J.; Matear, R.; Mikolajewicz, U.; Monfray, P.; Soldatov, V.; Spall, S. A.; Stouffer, R. (2004).

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The diagram on the right is a representation of how the comparative energetic effort for armor construction in diatoms, dinoflagellates and coccolithophores appear to operate. The

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are able to selectively choose prey on the basis of its size or shape and through chemical signals and may thus favor other prey that is available and not protected by coccoliths.

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Gafar, N. A., Eyre, B. D. and Schulz, K. G. (2019). "A comparison of species specific sensitivities to changing light and carbonate chemistry in calcifying marine phytoplankton".

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Irie, Takahiro; et al. (2010), "Increasing costs due to ocean acidification drives phytoplankton to be more heavily calcified: optimal growth strategy of coccolithophores",

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in calcium carbonate allows coccoliths to scatter more light than they absorb. This has two important consequences: 1) Surface waters become brighter, meaning they have a higher

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Buitenhuis, Erik T.; Pangerc, Tanja; Franklin, Daniel J.; Le Quéré, Corinne; Malin, Gill (2008), "Growth Rates of Six Coccolithoripd Strains as a Function of Temperature",

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Johns, Christopher T.; Grubb, Austin R.; Nissimov, Jozef I.; Natale, Frank; Knapp, Viki; Mui, Alwin; Fredricks, Helen F.; Van Mooy, Benjamin A. S.; Bidle, Kay D. (2019).

3879:"Detection of Phagotrophy in the Marine Phytoplankton Group of the Coccolithophores (Calcihaptophycidae, Haptophyta) During Nutrient‐replete and Phosphate‐limited Growth" 5548:

Sarmiento, Jorge L.; Hughes, Tertia M. C.; Stouffer, Ronald J.; Manabe, Syukuro (1998). "Simulated response of the ocean carbon cycle to anthropogenic climate warming".

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Young, J.R.; et al. (2009), "Coccolith function and morphogenesis: insights from appendage-bearing coccolithophores of the family syracosphaeraceae (haptophyta)",

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Mackinder; Wheeler, Glen; Schroeder, Declan; Riebesell, Ulf; Brownlee, Colin; et al. (2010), "Molecular Mechanisms Underlying Calcification in Coccolithophores",

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in the water column and a decrease in the vertical mixing of nutrients. However, a 2012 study estimated that the overall effect of coccolithophores on the increase in

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Linschooten, Cornelis; et al. (1991), "Role of the light-dark cycle and medium composition on the production of coccoliths by Emiliania huxleyi (haptophyceae)",

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Litchman, E.; et al. (2007), "The role of functional traits and trade-offs in structuring phytoplankton communities: scaling from cellular to ecosystem level",

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de Vries, Joost; Monteiro, Fanny; Wheeler, Glen; Poulton, Alex; Godrijan, Jelena; Cerino, Federica; Malinverno, Elisa; Langer, Gerald; Brownlee, Colin (2021-02-16).

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it is not clear whether global warming would result in net increase or decrease of coccolithophores. As they are calcifying organisms, it has been suggested that

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Charlson, Robert J.; Lovelock, James E.; Andreae, Meinrat O.; Warren, Stephen G. (1987). "Oceanic phytoplankton, atmospheric sulphur, cloud albedo and climate".

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Mayers, K.M.J.; Poulton, A.J.; Daniels, C.J.; Wells, S.R.; Woodward, E.M.S.; Tarran, G.A.; Widdicombe, C.E.; Mayor, D.J.; Atkinson, A.; Giering, S.L.C. (2019).

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Mayers, K.M.J.; Poulton, A.J.; Daniels, C.J.; Wells, S.R.; Woodward, E.M.S.; Tarran, G.A.; Widdicombe, C.E.; Mayor, D.J.; Atkinson, A.; Giering, S.L.C. (2019).

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Yunev, O.A.; et al. (2007), "Nutrient and phytoplankton trends on the western Black Sea shelf in response to cultural eutrophication and climate changes",

9444: 742:. These haploid cells can then divide further through mitosis or undergo sexual reproduction with other haploid cells. The resulting diploid cell goes through 722:(sexual) life cycle, and (c) coccolithophores tend to utilize a haplo-diplontic life cycle. Note that not all coccolithophores calcify in their haploid phase. 9493: 9380: 9375: 2457:

that was offered, rather than on their degree of calcification. In the same study, however, the authors found that predators which preyed on non-calcifying

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concentration results in an increase in calcification of these organisms. Decreasing coccolith mass is related to both the increasing concentrations of CO

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Kinkel, H.; et al. (2000), "Coccolithophores in the equatorial Atlantic Ocean: response to seasonal and Late Quaternary surface water variability",

2963: 2059: 6200: 9576: 9488: 9395: 8959: 6471:"Emiliania huxleyi increases calcification but not expression of calcification-related genes in long-term exposure to elevated temperature and p CO 2" 9385: 9313: 9297: 9214: 9204: 9170: 8997: 4836:

Henderiks, Jorijntje (2008). "Coccolithophore size rules — Reconstructing ancient cell geometry and cellular calcite quota from fossil coccoliths".

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Young, J. R. (1994). "Functions of coccoliths", in Coccolithophores, eds A. Winter and W. G. Siesser (Cambridge: Cambridge University Press), 63–82.

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Smith, H.E.K.; et al. (2012), "Predominance of heavily calcified coccolithophores at low CaCO3 saturation during winter in the Bay of Biscay",

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Bates; Michaels, Anthony F.; Knap, Anthony H.; et al. (1996), "Alkalinity changes in the Sargasso Sea; geochemical evidence of calfication?",

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Evolutionary history of coccolithophores: (A) Coccolithophore species richness over time; (B) The fossil record of major coccolithophore

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did not differ significantly from those on similar sized non-calcifying phytoplankton. In laboratory experiments the heterotrophic dinoflagellate

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de Vargas, C.; Aubrey, M.P.; Probert, I.; Young, J. (2007). "From coastal hunters to oceanic farmers.". In Falkowski, P.G.; Knoll, A.H. (eds.).

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are monitoring the responses of coccolithophore populations to varying pH's and working to determine environmentally sound measures of control.

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Coccolithophores (or coccolithophorids, from the adjective) form a group of about 200 phytoplankton species. They belong either to the kingdom

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Billard, Chantal; Inouye, Isoa (August 17, 2004). "What is new in coccolithophore biology?". In Thierstein, Hans R.; Young, Jeremy R. (eds.).

9753: 9527: 9118: 686:, coils and uncoils in response to environmental stimuli. Although poorly understood, it has been proposed to be involved in prey capture. 5111:

Geisen, M.; et al. (August 17, 2004). "Species level variation in coccolithophores=". In Thierstein, Hans R.; Young, Jeremy R. (eds.).

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is a region of elevated summertime upper ocean calcite concentration derived from coccolithophores, despite the region being known for its

6551:"Nannofossil carbonate fluxes during the Early Cretaceous: Phytoplankton response to nutrification episodes, atmospheric CO2, and anoxia" 5747: 3011: 2380:

Representation of comparative energetic effort for armor construction in three major shell-forming phytoplankton taxa as a function of

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depths. These coccolithophores increase in abundance when the nutricline and thermocline are deep and decrease when they are shallow.

3174: 2183:, and of other similar rocks in many other parts of the world. At the present day sedimented coccoliths are a major component of the 7100:"Grazing in the heterotrophic dinoflagellate Oxyrrhis marina: Size selectivity and preference for calcified Emiliania huxleyi cells" 6469:

Benner, Ina; Diner, Rachel E.; Lefebvre, Stephane C.; Li, Dian; Komada, Tomoko; Carpenter, Edward J.; Stillman, Jonathon H. (2013).

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Vardi, A.; et al. (2012), "Host–virus dynamics and subcellular controls of cell fate in a natural coccolithophore population",

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in the ocean. Finally, field evidence of coccolithophore fossils in rock were used to show that the deep-sea fossil record bears a

2111:. Calcium carbonate is transparent, so the organisms' photosynthetic activity is not compromised by encapsulation in a coccosphere. 9688: 4588:

Calbet, Albert; Landry, Michael R. (2004). "Phytoplankton growth, microzooplankton grazing, and carbon cycling in marine systems".

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in the world's oceans. This lower calcification is assumed to put coccolithophores at ecological disadvantage. Some species like

9727: 7941: 4748:"The role of dissolved infochemicals in mediating predator-prey interactions in the heterotrophic dinoflagellate Oxyrrhis marina" 2419:, while others found high microzooplankton grazing rates on natural coccolithophore communities. In 2020, researchers found that 2052: 1257:

evolutionary framework, but instead a "Cheshire Cat" ecological dynamic. More recent work has suggested that viral synthesis of

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associated with these faster growth rates include a smaller cell radius and lower cell volume than other types of phytoplankton.

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phase, lack radial symmetry, and are composed of anywhere from hundreds to thousands of similar minute (ca 0.1 μm) rhombic

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due to decreasing global temperatures, with species that produced large and heavily calcified coccoliths most heavily affected.

8465: 7856: 3183: 1305:, when more than 90% of coccolithophore species became extinct. Coccoliths reached another, lower apex of diversity during the 7564: 6593:

Erba, Elisabetta (2006). "The first 150 million years history of calcareous nannoplankton: Biosphere–geosphere interactions".

4075:"Association of sinking organic matter with various types of mineral ballast in the deep sea: Implications for the rain ratio" 2368:

as their source of structural elements in the form of cellulose should be facilitated by the ocean acidification-associated CO

2230:(A) Transport processes include the transport into the cell from the surrounding seawater of primary calcification substrates 7693: 5870: 4224: 3980: 3947: 3854: 3430: 1302: 796:

species are located, the ocean currents also can determine the location where certain species of coccolithophores are found.

9732: 6040: 2437:, which was hypothesised to be due to size selective feeding behaviour, since calcified cells are larger than non-calcified 1083:

Coccolithophores are one of the more abundant primary producers in the ocean. As such, they are a large contributor to the

1306: 5458:"The influence of environmental variability on the biogeography of coccolithophores and diatoms in the Great Calcite Belt" 3930:

Bown, Paul R.; Lees, Jackie A.; Young, Jeremy R. (2004). "Calcareous nannoplankton evolution and diversity through time".

6686:

Van De Waal, Dedmer B.; John, Uwe; Ziveri, Patrizia; Reichart, Gert-Jan; Hoins, Mirja; Sluijs, Appy; Rost, Björn (2013).

2376: 939:, but they are heavily calcified and make important contributions to global calcification. Unmarked scale bars 5 μm. 5509:"Calcium carbonate measurements in the surface global ocean based on Moderate-Resolution Imaging Spectroradiometer data" 8578: 7820: 6637:"A unifying concept of coccolithophore sensitivity to changing carbonate chemistry embedded in an ecological framework" 4960: 3091:

However, the production of calcium carbonate drives surface alkalinity down, and in conditions of low alkalinity the CO

2045: 727: 7565:"Calcareous Nannofossil Assemblage Changes Across the Paleocene-Eocene Thermal Maximum: Evidence from a Shelf Setting" 7467: 3168:

might be (study results are mixed). Also, highly calcified coccolithophorids have been found in conditions of low CaCO

2461:

grew faster than those fed with calcified cells. In 2018, Strom et al. compared predation rates of the dinoflagellate

553:

precipitation of calcium carbonate during coccolith formation reduces the total alkalinity of seawater and releases CO

6150: 5876: 5120: 4923: 4439: 4239:

Young, J. R. (1987). Possible Functional Interpretations of Coccolith Morphology. New York: Springer-Verlag, 305–313.

3916: 3261: 7391:

Beaufort, L.; et al. (2011), "Sensitivity of coccolithophores to carbonate chemistry and ocean acidification",

4207:

Rost, Björn; Riebesell, Ulf (2004). "Coccolithophores and the biological pump: Responses to environmental changes".

3413:

Rost, Björn; Riebesell, Ulf (2004). "Coccolithophores and the biological pump: Responses to environmental changes".

9789: 8615: 3364:"Influence of the Calcium Carbonate Shell of Coccolithophores on Ingestion and Growth of a Dinoflagellate Predator" 375:-related coccolithophore blooms, as these blooms lead to a decrease in nutrient flow to lower levels of the ocean. 17: 9758: 5386:

Daniels, C.J., Sheward, R.M. and Poulton, A.J. (2014) "Biogeochemical implications of comparative growth rates of

2331:. Reduced intra-cellular pH would severely affect the entire cellular machinery and require other processes (e.g. 8610: 6048: 2449:

as well as calcified strains that differed in the degree of calcification. They found that the ingestion rate of

600: 7597:

Lloyd, G.T.; et al. (2011), "Quantifying the deep-sea rock and fossil record bias using coccolithophores",

6434:

Taylor, Alison R.; Brownlee, Colin; Wheeler, Glen L. (2012). "Proton channels in algae: Reasons to be excited".

460:, according to the newer biological classification system. Within the Hacrobia, the coccolithophores are in the 3249:. They are the largest global source of biogenic calcium carbonate, and significantly contribute to the global 3004: 3290:

The coccolithophorids help in regulating the temperature of the oceans. They thrive in warm seas and release

2195: 2144:

control the shape and growth of these crystals. As each scale is produced, it is exported in a Golgi-derived

730:, and is characterized by an alternation of both asexual and sexual phases. The asexual phase is known as the 280:, according to a newer biological classification system. Within the Hacrobia, the coccolithophores are in the 9740: 8694: 8684: 8667: 3237: 2867: 1922: 1917: 1750: 1551: 1120:

of waters and thus, coccolithophore blooms in these high nitrogen and phosphorus, low silicate environments.

142: 3803:"A review of selected aspects of coccolithophore biology with implications for paleobiodiversity estimation" 347:

Coccolithophores are ecologically important, and biogeochemically they play significant roles in the marine

8530: 6635:

Bach, Lennart Thomas; Riebesell, Ulf; Gutowska, Magdalena A.; Federwisch, Luisa; Schulz, Kai Georg (2015).

2907: 2273:

crystals. The completed coccolith (gray plate) is a complex structure of intricately arranged CAPs and CaCO

994: 449: 269: 7007:"Growth and mortality of coccolithophores during spring in a temperate Shelf Sea (Celtic Sea, April 2015)" 5097:. Washington, D.C.: Environmental Information Coalition, National Council for Science and the Environment. 4643:"Growth and mortality of coccolithophores during spring in a temperate Shelf Sea (Celtic Sea, April 2015)" 616:

Coccolithophores are spherical cells about 5–100 micrometres across, enclosed by calcareous plates called

8455: 7966: 6110:

Gardin, Silvia; Krystyn, Leopold; Richoz, Sylvain; Bartolini, Annachiara; Galbrun, Bruno (October 2012).

5234:

Okada; Honjo, Susumu; et al. (1973), "The distribution of oceanic coccolithophores in the Pacific",

2596: 1745: 416:. It is also the fastest growing coccolithophore in laboratory cultures. It is studied for the extensive 6201:"Microscopic marine plants bioengineer their environment to enhance their own growth - The Conversation" 2496: 8445: 7946: 7911: 7792: 6111: 6057: 5911:

Houdan, A.; et al. (2004), "Toxicity of coastal coccolithophores (Prymnesiophyceae, Haptophyta)",

3318: 3034: 2958: 2690: 2565: 2032: 1979: 1061: 5592: 4557:"The Calcium Carbonate Shell of Emiliania huxleyi Provides Limited Protection Against Viral Infection" 2706: 2472:

calcification protects coccolithophores from microzooplankton predation could not be fully clarified.

9794: 9653: 8500: 7487: 7182:"Phytoplankton defenses: Do Emiliania huxleyi coccoliths protect against microzooplankton predators?" 4160:"Marine calcification as a source of carbon dioxide: Positive feedback of increasing atmospheric CO2" 3362:

Haunost, Mathias; Riebesell, Ulf; D'Amore, Francesco; Kelting, Ole; Bach, Lennart T. (30 June 2021).

3264:

55 million years ago. This period is thought to correspond most directly to the current levels of CO

3125:

in to constantly pump H ions out of the cell during coccolith production. This allows them to avoid

2997: 2922: 2695: 1973: 1632: 1362: 1297:

boundary. Diversity steadily increased over the course of the Mesozoic, reaching its apex during the

1254: 1084: 356: 5853: 2199:

Energetic costs of coccolithophore calcification. Energetic costs reported as a percentage of total

933:

Larger coccolithophores such as the species above are less numerous than the smaller but ubiquitous

8162: 8034: 8021: 7783:

University of California, Berkeley. Museum of Paleontology: "Introduction to the Prymnesiophyta".

6214:

Westbroek, P.; et al. (1983), "Calcification in Coccolithophoridae: Wasteful or Functional?",

2244:(black arrows) and the removal of the end product H from the cell (gray arrow). The transport of Ca 1519: 6814:

Reinfelder, John R. (2011). "Carbon Concentrating Mechanisms in Eukaryotic Marine Phytoplankton".

4978:

Young, J.R.; Karen, H. (2003). "Biomineralization Within Vesicles: The Calcite of Coccoliths". In

3760:

Bentaleb, I.; et al. (1999), "Silicate as regulating nutrient in phytoplankton competition",

8699: 8440: 8420: 7886: 3207: 3118: 2943: 2912: 2589: 2226:

production. The diagram on the right shows the energetic costs of coccolithophore calcification:

1755: 1627: 1386: 1087:

of the tropical and subtropical oceans, however, exactly how much has yet to have been recorded.

877:

environments, unique oceanic topography, and pockets of isolated high or low water temperatures.

433: 249: 2294: 9837: 9615: 8571: 8450: 8302: 7813: 5848: 3332: 2917: 2790: 1944: 1769: 1141: 562: 541:

sediments. Their calcareous shell increases the sinking velocity of photosynthetically fixed CO

6112:"Where and when the earliest coccolithophores?: Where and when the earliest coccolithophores?" 9822: 8662: 8495: 8322: 8265: 8000: 7866: 7782: 6859:"Coccolith arrangement follows Eulerian mathematics in the coccolithophore Emiliania huxleyi" 4703:

Tillmann, Urban (2004). "Interactions between Planktonic Microalgae and Protozoan Grazers1".

3910: 3697:

Egge, JK; Aksnes, DL (1992), "Silicate as regulating nutrient in phytoplankton competition",

3254: 3223: 3190: 3121:

of coccolithophore species over longer periods of time. For example, coccolithophores use H

2311: 2180: 1507: 1380: 1368: 1262: 1212:

Coccolithophorids are predominantly found as single, free-floating haploid or diploid cells.

660: 9781: 9771: 9719: 3736: 9832: 9817: 9675: 8247: 8047: 7951: 7916: 7718: 7648: 7606: 7572: 7442: 7326: 7284: 7229:

Mejia, R. (2011), "Will Ion Channels Help Coccolithophores Adapt to Ocean Acidification?",

7193: 7018: 6979: 6935: 6823: 6758: 6699: 6648: 6602: 6562: 6383:

Taylor, Alison R.; Chrachri, Abdul; Wheeler, Glen; Goddard, Helen; Brownlee, Colin (2011).

6342: 6300: 6238: 6061: 5994: 5948: 5704: 5661: 5607: 5557: 5520: 5469: 5280: 5243: 5184: 5009: 4845: 4792: 4654: 4597: 4511: 4342: 4271: 4171: 4132: 4086: 4045: 4006: 3817: 3706: 3662: 3585: 3528: 3462: 2847: 2611: 2179:, a Late Cretaceous rock formation which outcrops widely in southern England and forms the 1539: 1321: 770: 759: 352: 7180:

Strom, Suzanne L.; Bright, Kelley J.; Fredrickson, Kerri A.; Cooney, Elizabeth C. (2018).

5981:

Frada, M.; et al. (2008), "The "Cheshire Cat" escape strategy of the coccolithophore

3164:, however, are not affected in this way, while the most abundant coccolithophore species, 3080:

Because coccolithophores are photosynthetic organisms, they are able to use some of the CO

2393: 1111:

within phytoplankton communities. Each ratio essentially tips the odds in favor of either

8: 8830: 8490: 8425: 8192: 8181: 8058: 8041: 7683: 6835: 6073: 3737:"Life at the Edge of Sight — Scott Chimileski, Roberto Kolter | Harvard University Press" 3110: 2931: 2852: 2770: 2401: 2307: 2219: 2184: 1461: 1108: 1020: 699: 558: 417: 364: 323: 273: 7722: 7652: 7610: 7576: 7446: 7330: 7288: 7197: 7022: 6983: 6939: 6827: 6762: 6703: 6652: 6606: 6566: 6346: 6304: 6242: 6065: 5998: 5952: 5708: 5665: 5611: 5561: 5524: 5473: 5284: 5247: 5188: 5013: 4952: 4849: 4796: 4658: 4601: 4515: 4455:

Brussaard, Corina P. D. (2004). "Viral Control of Phytoplankton Populations-a Review1".

4346: 4275: 4175: 4136: 4090: 4049: 4010: 3878: 3821: 3710: 3666: 3589: 3532: 3466: 389: 8679: 8657: 8645: 8620: 7734: 7664: 7622: 7540: 7513: 7416: 7300: 7253: 7211: 7036: 6951: 6895: 6858: 6722: 6687: 6495: 6470: 6411: 6384: 6261: 6183: 6017: 5728: 5633: 5573: 5370: 5157: 5032: 4818: 4728: 4716: 4672: 4623: 4532: 4499: 4480: 4468: 4431: 4363: 4330: 4104: 3680: 3551: 3488: 2936: 2872: 2754: 2324: 1994: 1984: 1835: 1801: 1718: 1711: 1622: 1471: 1072: 978: 972: 964: 340:

remain elusive. One key function may be that the coccosphere offers protection against

137: 7367: 6991: 5773: 5292: 3773: 9766: 9662: 9555: 9550: 8969: 8689: 8652: 8640: 8630: 8564: 8385: 8380: 8375: 8197: 8116: 7961: 7881: 7806: 7738: 7689: 7626: 7545: 7472: 7433:

Tyrell, T.; et al. (1999), "Optical impacts of oceanic coccolithophore blooms",

7408: 7371: 7338: 7258: 7040: 6900: 6882: 6839: 6727: 6500: 6451: 6416: 6266: 6179: 6146: 6127: 6085: 6077: 6022: 5964: 5960: 5866: 5720: 5507:

Balch, W. M.; Gordon, Howard R.; Bowler, B. C.; Drapeau, D. T.; Booth, E. S. (2005).

5362: 5358: 5255: 5116: 5037: 4956: 4919: 4861: 4810: 4720: 4676: 4537: 4472: 4435: 4368: 4220: 3976: 3943: 3898: 3850: 3802: 3556: 3492: 3480: 3426: 3385: 3291: 3038: 2984: 2877: 2658: 2584: 2415: 2258: 2211: 2145: 2125: 2120: 1999: 1909: 1578: 1512: 1437: 1374: 1345: 1328: 1145: 1067: 935: 861: 837: 813: 763: 507: 490: 480: 476: 465: 453: 380: 327: 296: 285: 7514:"A voltage-gated H channel underlying pH homeostasis in calcifying coccolithophores" 7304: 7215: 6955: 6385:"A Voltage-Gated H Channel Underlying pH Homeostasis in Calcifying Coccolithophores" 6187: 5637: 5374: 4732: 4627: 4484: 4108: 3684: 3243: 851: 264:

community. They form a group of about 200 species, and belong either to the kingdom

9537: 9521: 9358: 9255: 9025: 9007: 8954: 8949: 8812: 8763: 8604: 8327: 8307: 8231: 8105: 8052: 7991: 7726: 7709:

Larsen, S. H. (2005). "Solar variability, dimethyl sulphide, clouds, and climate".

7668: 7656: 7614: 7584: 7580: 7535: 7525: 7450: 7420: 7400: 7363: 7334: 7292: 7248: 7238: 7201: 7157: 7147: 7111: 7078: 7068: 7026: 6987: 6943: 6908: 6890: 6872: 6831: 6796: 6766: 6717: 6707: 6656: 6610: 6570: 6528: 6490: 6482: 6443: 6406: 6396: 6350: 6308: 6256: 6246: 6175: 6123: 6069: 6012: 6002: 5956: 5920: 5858: 5732: 5712: 5669: 5623: 5615: 5577: 5565: 5528: 5477: 5429: 5403: 5354: 5313: 5288: 5251: 5192: 5161: 5149: 5069: 5027: 5017: 4979: 4948: 4894: 4857: 4853: 4822: 4800: 4759: 4712: 4662: 4613: 4605: 4568: 4527: 4519: 4464: 4427: 4399: 4358: 4350: 4289: 4279: 4212: 4189: 4179: 4140: 4094: 4053: 4014: 3968: 3935: 3890: 3825: 3769: 3714: 3670: 3593: 3546: 3536: 3470: 3418: 3375: 3313: 2579: 2538: 1959: 1949: 1929: 1546: 1466: 574: 546: 472: 413: 341: 292: 197: 6447: 5689: 5140:

Jordan, R. W.; Chamberlain, A.H.L. (1997), "Biodiversity among haptophyte algae",

4034:"Ratio of coccolith CaCO3to foraminifera CaCO3in late Holocene deep sea sediments" 2506: 1023:

due to increasing carbon dioxide could severely affect coccolithophores. Recent CO

234:

Coccolithophore cells are covered with protective calcified (chalk) scales called

9516: 9017: 8992: 8979: 8941: 8856: 8839: 8804: 8435: 8312: 8138: 8122: 7926: 7851: 7530: 7243: 7031: 7006: 6747:"Evolutionary and ecological perspectives on carbon acquisition in phytoplankton" 6712: 6688:"Ocean Acidification Reduces Growth and Calcification in a Marine Dinoflagellate" 6661: 6636: 6614: 6401: 6313: 6288: 6251: 4693:, Eds A. Winter and W. G. Siesser (Cambridge: Cambridge University Press), 63–82. 4667: 4642: 4144: 3972: 3939: 3844: 3323: 3308: 3285: 2902: 2800: 2759: 2749: 2429: 2365: 2141: 1825: 1701: 1615: 1583: 1298: 1289:. The oldest known coccolithophores are known from the Late Triassic, around the 1137: 1056: 801: 656: 530: 348: 9745: 9667: 6532: 5862: 4898: 4216: 3829: 3787: 3634:"Biogeography and dispersal of micro-organisms: a review emphasizing protists", 3422: 1337: 1002: 738:

phase. During the haploid phase, coccolithophores produce haploid cells through

493:. However, there are Prymnesiophyceae species lacking coccoliths (e.g. in genus 344:

predation, which is one of the main causes of phytoplankton death in the ocean.

9638: 9508: 9327: 9188: 9149: 9002: 8900: 8848: 8635: 8505: 8236: 8151: 8027: 7769: 7761:– illustrated guide to the taxonomy of coccolithophores and other nannofossils. 6673: 6367: 5494: 5443: 5327: 5317: 4885: 4746:

Breckels, M. N.; Roberts, E. C.; Archer, S. D.; Malin, G.; Steinke, M. (2011).

4018: 3597: 3503: 3400: 3085: 3042: 2862: 2842: 2810: 2715: 2647: 2557: 2543: 2352: 2332: 2320: 2200: 1967: 1830: 1817: 1671: 1644: 1639: 1117: 982: 752: 707: 582: 385: 372: 360: 304: 300: 212: 167: 7296: 7073: 7056: 6947: 6800: 6771: 6746: 6355: 6330: 5834:"Coccolithophores and the biological pump: responses to environmental changes" 5196: 5153: 4609: 4573: 4556: 4184: 4159: 3675: 3380: 3363: 2222:. Coccolithophores are the major planktonic group responsible for pelagic CaCO 782: 9811: 9470: 9449: 8932: 8892: 8736: 8515: 8485: 8370: 8365: 8092: 8069: 8006: 7976: 7971: 6886: 6283:

Krumhardt, Kristen M.; Lovenduski, Nicole S.; Iglesias-Rodriguez, M. Debora;

6081: 4865: 4814: 4404: 4387: 3902: 3484: 3389: 3273: 3130: 2815: 2669: 2601: 2514: 2262: 2207: 2133: 2073: 1989: 1934: 1774: 1737: 1600: 1558: 1529: 1524: 1408: 1258: 1221: 1040: 998: 586: 503: 429: 253: 219: 76: 7152: 7135: 6282: 6007: 5925: 5716: 5482: 5457: 5407: 5022: 4805: 4780: 4764: 4747: 4523: 3541: 3475: 3450: 1027:

increases have seen a sharp increase in the population of coccolithophores.

518:) which cover the cell surface in the form of a spherical coating, called a 9282: 8883: 8510: 8405: 8332: 8287: 8259: 8156: 8132: 7981: 7906: 7876: 7871: 7861: 7549: 7512:

Taylor, A.R.; Chrachri, A.; Wheeler, G.; Goddard, H.; Brownlee, C. (2011).

7412: 7375: 7262: 6904: 6843: 6731: 6504: 6486: 6455: 6420: 6284: 6270: 6089: 6026: 5968: 5724: 5366: 5041: 4724: 4541: 4476: 4372: 4354: 3560: 3250: 3122: 3030: 2857: 2805: 2795: 2735: 2631: 2606: 2533: 2528: 2189: 2004: 1681: 1676: 1658: 1476: 1282: 1076: 1035: 792: 672: 652: 648: 640: 625: 621: 37: 6912: 5833: 5796: 4500:"The mutual interplay between calcification and coccolithovirus infection" 2310:

is presently unknown. Cell physiological examinations found the essential

919: 9701: 9647: 9344: 9158: 9080: 8918: 8874: 8724: 8475: 8460: 8410: 8275: 8099: 7896: 7891: 7846: 7730: 7454: 7354:

Marsh, M.E. (2003), "Regulation of CaCO3 formation in coccolithophores",

6575: 6550: 5673: 5619: 5533: 5508: 4099: 4074: 4058: 4033: 3963:

Hay, William W. (2004). "Carbonate fluxes and calcareous nannoplankton".

3246: 3046: 2953: 2948: 2897: 2837: 2829: 2740: 2519: 2463: 2176: 2092: 1954: 1939: 1882: 1796: 1563: 1495: 1351: 1266: 1253:" between coccolithophores and these viruses does not follow the classic 1242: 1044: 910: 889: 845: 683: 519: 421: 420:

it forms in nutrient depleted waters after the reformation of the summer

397: 368: 332: 312: 257: 128: 51: 7404: 5433: 4618: 4294: 1445: 903: 620:, which are about 2–25 micrometres across. Each cell contains two brown 119: 9336: 9273: 8964: 8910: 8360: 8297: 8226: 8218: 8016: 8011: 7956: 6877: 4986:. Washington, D.C.: Mineralogical Society of America. pp. 189–216. 4422:

Hamm, Christian; Smetacek, Victor (2007). "Armor: Why, when, and How".

4193: 3719: 3295: 3134: 2663: 2574: 2287: 2283: 2266: 2088: 1534: 1278: 1269:

arms race at least between the coccolithoviruses and diploid organism.

1100: 841: 636: 617: 523: 495: 468: 409: 355:. Depending on habitat, they can produce up to 40 percent of the local 288: 187: 96: 61: 9706: 8556: 7206: 7181: 7116: 7099: 7083: 7057:"The Possession of Coccoliths Fails to Deter Microzooplankton Grazers" 6588: 6586: 6544: 6542: 6475:

Philosophical Transactions of the Royal Society B: Biological Sciences

6378: 6376: 5074: 4417: 4415: 4284: 4259: 4158:

Frankignoulle, Michel; Canon, Christine; Gattuso, Jean-Pierre (1994).

3894: 2441:. In 2015, Harvey et al. investigated predation by the dinoflagellate 9180: 8777: 8715: 8587: 8430: 8415: 8355: 8337: 8317: 8292: 8253: 8241: 8081: 7660: 7618: 7162: 5816: 5628: 2397: 2381: 2340: 2249: 2077: 1877: 1870: 1764: 1610: 1568: 1310: 1250: 1238: 1226: 870: 719: 711: 679: 676: 605: 515: 485: 393: 308: 235: 154: 101: 9693: 9609: 3272:

similar to the one that is widely accepted to affect the land-based

9632: 9140: 8795: 8786: 8674: 8591: 8480: 8186: 8145: 8127: 7901: 7838: 7829: 6583: 6539: 6373: 4412: 3417:. Berlin, Heidelberg: Springer Berlin Heidelberg. pp. 99–125. 3269: 3173:

from future research in this area. Groups like the European-based

3126: 2679: 2488: 2458: 2389: 1865: 1855: 1590: 1573: 1294: 1286: 1104: 1096: 664: 632: 550: 534: 457: 445: 425: 401: 277: 265: 261: 230: 177: 91: 86: 71: 66: 56: 7274: 6928:

Journal of the Marine Biological Association of the United Kingdom

5569: 4781:"Covariation of metabolic rates and cell size in coccolithophores" 3877:

Avrahami, Yoav; Frada, Miguel J. (31 March 2020). Mock, T. (ed.).

3616:

Transactions of the Gulf Coast Association of Geological Societies

3361: 2339:

perturbation events even though evolutionary adaption to changing

1013: 9051: 8540: 8470: 8347: 8086: 7758: 7134:

Harvey, Elizabeth L.; Bidle, Kay D.; Johnson, Matthew D. (2015).

6672:

Material was copied from this source, which is available under a

6366:

Material was copied from this source, which is available under a

5493:

Material was copied from this source, which is available under a

5442:

Material was copied from this source, which is available under a

5326:

Material was copied from this source, which is available under a

5093:

Hogan, M.C. ""Coccolithophores"". In Cleveland, Cutler J. (ed.).

3652: 3502:

Material was copied from this source, which is available under a

3399:

Material was copied from this source, which is available under a

2361: 2150: 2129: 1860: 1785: 1706: 1595: 1481: 1129: 881:

same values in between that of the lower and upper photic zones.

874: 748: 743: 739: 735: 731: 668: 667:

structures, which are involved not only in motility, but also in

644: 589: 578: 499:), so not every member of Prymnesiophyceae is a coccolithophore. 337: 106: 81: 6039:

Taylor, Alison R.; Brownlee, Colin; Wheeler, Glen (2017-01-03).

4121: 3104: 1188:

were shown to be non-toxic as were species of the coastal genus

1051: 367:

increases, their coccoliths may become even more important as a

8202: 8173: 6634: 3575: 2674: 2636: 2348: 2107:

The primary constituent of coccoliths is calcium carbonate, or

1889: 1290: 1133: 1112: 986: 715: 461: 281: 7053: 6289:"Coccolithophore growth and calcification in a changing ocean" 4388:"On the Genesis and Function of Coccolithophore Calcification" 3448: 3253:. They are the main constituent of chalk deposits such as the 3029:

Coccolithophores have both long and short term effects on the

2087:

Each coccolithophore encloses itself in a protective shell of

9680: 7638: 6863: 6685: 6668: 6362: 6331:"A global compilation of coccolithophore calcification rates" 5650: 5547: 5489: 5438: 5322: 3498: 3395: 2252:

to the CV is the dominant cost associated with calcification.

2238: 2231: 2108: 1605: 855:

Size comparison between the relatively large coccolithophore

316: 7511: 7179: 6382: 6109: 5748:"What's fueling the rise of coccolithophores in the oceans?" 8520: 7798: 5213: 5173: 4555:

Haunost, Mathias; Riebesell, Ulf; Bach, Lennart T. (2020).

2364:

may even profit from chemical changes since photosynthetic

1723: 1662: 359:. They are of particular interest to those studying global 6972:

Deep Sea Research Part II: Topical Studies in Oceanography

6327: 5113:

Coccolithophores-from molecular processes to global impact

4916:

Coccolithophores-from molecular processes to global impact

4745: 3999:

Deep Sea Research Part II: Topical Studies in Oceanography

3451:"Haplo-diplontic life cycle expands coccolithophore niche" 526:, and are able to photosynthesise as well as ingest prey. 7562: 7136:"Consequences of strain variability and calcification in 7004: 6856: 5590: 5455: 5058: 4640: 4260:"Environmental controls on coccolithophore calcification" 4157: 2347:

Silicate- or cellulose-armored functional groups such as

2344:

chemistry perturbations can be compensated by evolution.

1010:) alongside the North Atlantic and North Pacific oceans. 726:

The complex life cycle of coccolithophores is known as a

7765:

INA — International Nannoplankton Association

7491: 6925: 5506: 4497: 4327: 3995: 859:

and the relatively small but ubiquitous coccolithophore

647:

are located on either side of the cell and surround the

27:

Unicellular algae responsible for the formation of chalk

7764: 5177:

Deep-Sea Research Part I: Oceanographic Research Papers

4881: 2328: 2261:

include the synthesis of CAPs (gray rectangles) by the

2164: 755:

may affect the frequency with which each phase occurs.

557:. Thus, coccolithophores play an important role in the 6674: