Coccolithophore
Coccolithophore Temporal range:
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Coccolithus pelagicus | |
Scientific classification | |
Domain: | Eukaryota |
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(unranked): | Haptophyta |
Class: | |
Order: |
![](http://upload.wikimedia.org/wikipedia/commons/thumb/e/e3/Coccolithophores.png/240px-Coccolithophores.png)
Coccolithophores, or coccolithophorids, are
.Coccolithophores are the most productive
Coccolithophores are ecologically important, and biogeochemically they play significant roles in the marine biological pump and the carbon cycle.[2][1] Depending on habitat, they can produce up to 40 percent of the local marine primary production.[3] They are of particular interest to those studying global climate change because, as ocean acidity increases, their coccoliths may become even more important as a carbon sink.[4] Management strategies are being employed to prevent eutrophication-related coccolithophore blooms, as these blooms lead to a decrease in nutrient flow to lower levels of the ocean.[5]
The most abundant species of coccolithophore,
Overview
Coccolithophores (or coccolithophorids, from the adjective
Coccolithophores are single-celled
As of 2021, it is not known why coccolithophores calcify and how their ability to produce coccoliths is associated with their ecological success.
Structure
![](http://upload.wikimedia.org/wikipedia/commons/thumb/2/23/Diagram_of_a_coccolithophore_cell_and_its_shield_of_coccoliths.png/240px-Diagram_of_a_coccolithophore_cell_and_its_shield_of_coccoliths.png)
Coccolithophores are spherical cells about 5–100 micrometres across, enclosed by calcareous plates called
Enclosed in each coccosphere is a single cell with
Ecology
Life history strategy
The complex life cycle of coccolithophores is known as a
Coccolithophores reproduce asexually through binary fission. 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.[44]
K or r- selected strategies 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.[44] Most 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.[45] 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.[44]
Global distribution
![](http://upload.wikimedia.org/wikipedia/commons/thumb/b/b8/Coccolithophore%2BAbundance.png/370px-Coccolithophore%2BAbundance.png)
Coccolithophores occur throughout the world's oceans. Their distribution varies vertically by stratified layers in the ocean and geographically by different temporal zones.
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
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.[50] The highest diversity of coccolithophores in the Pacific Ocean was in an area of the ocean considered the Central North Zone which is an area between 30 oN and 5 oN, 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.[50]
In the Atlantic Ocean, the most abundant species are
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 ocean's properties, such as coastal and equatorial
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 same values in between that of the lower and upper photic zones.[47]
Great Calcite Belt
The Great Calcite Belt of the Southern Ocean is a region of elevated summertime upper ocean calcite concentration derived from coccolithophores, despite the region being known for its diatom 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.[56]
The Great Calcite Belt, defined as an elevated particulate inorganic carbon (PIC) feature occurring alongside seasonally elevated chlorophyll a in austral spring and summer in the Southern Ocean,[57] plays an important role in climate fluctuations,[58][59] accounting for over 60% of the Southern Ocean area (30–60° S).[60] The region between 30° and 50° S has the highest uptake of anthropogenic carbon dioxide (CO2) alongside the North Atlantic and North Pacific oceans.[61]
Effect of global climate change on distribution
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 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 ocean acidification due to increasing carbon dioxide could severely affect coccolithophores.[51] Recent CO2 increases have seen a sharp increase in the population of coccolithophores.[62]
Role in the food web
![](http://upload.wikimedia.org/wikipedia/commons/thumb/8/83/Bloom_in_the_Barents_Sea.jpg/310px-Bloom_in_the_Barents_Sea.jpg)
Coccolithophores are one of the more abundant primary producers in the ocean. As such, they are a large contributor to the primary productivity of the tropical and subtropical oceans, however, exactly how much has yet to have been recorded.[66]
Dependence on nutrients
The ratio between the concentrations of nitrogen, phosphorus and silicate in particular areas of the ocean dictates competitive dominance within phytoplankton communities. Each ratio essentially tips the odds in favor of either diatoms 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 agricultural processes lead to eutrophication of waters and thus, coccolithophore blooms in these high nitrogen and phosphorus, low silicate environments.[5]
Impact on water column productivity
The calcite 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 albedo, and 2) there is induced photoinhibition, 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 stratification 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 radiative forcing of the ocean is less than that from anthropogenic factors.[67] Therefore, the overall result of large blooms of coccolithophores is a decrease in water column productivity, rather than a contribution to global warming.
Predator-prey interactions
Their predators include the common predators of all phytoplankton including small fish, zooplankton, and shellfish larvae.[45][68] Viruses specific to this species have been isolated from several locations worldwide and appear to play a major role in spring bloom dynamics.
Toxicity
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 Prymnesium Massart and Chrysochromulina Lackey. Members of the genus Prymnesium 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 coccolithophore genera Emiliania, Gephyrocapsa, Calcidiscus and Coccolithus were shown to be non-toxic as were species of the coastal genus Hymenomonas, however several species of Pleurochrysis and Jomonlithus, both coastal genera were toxic to Artemia.[68]
Community interactions
Coccolithophorids are predominantly found as single, free-floating haploid or diploid cells.[46]
Competition
Most phytoplankton 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 (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.[69] Trade-offs associated with these faster growth rates include a smaller cell radius and lower cell volume than other types of phytoplankton.
Viral infection and coevolution
Giant
Evolution and diversity
Coccolithophores are members of the clade
![](http://upload.wikimedia.org/wikipedia/commons/thumb/6/69/Evolutionary_history_of_coccolithophores.jpg/370px-Evolutionary_history_of_coccolithophores.jpg)
Coccolithophore shells
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- Exoskeleton: coccospheres and coccoliths
Each coccolithophore encloses itself in a protective shell of
Composition
The primary constituent of coccoliths is calcium carbonate, or chalk. Calcium carbonate is transparent, so the organisms' photosynthetic activity is not compromised by encapsulation in a coccosphere.[45]
Formation
Coccoliths are produced by a
Function
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 pH. 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.[75] 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.[76] More specific, defensive properties of coccoliths may include protection from osmotic changes, chemical or mechanical shock, and short-wavelength light.[41] 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.[77] Coccolith appendages have also been proposed to serve several functions, such as inhibiting grazing by zooplankton.[53]
Uses
Coccoliths are the main component of
- (A) Transport processes include the transport into the cell from the surrounding seawater of primary calcification substrates
- (B) Golgi complex (white rectangles) that regulate the nucleation and geometry of CaCO3 crystals. The completed coccolith (gray plate) is a complex structure of intricately arranged CAPs and CaCO3 crystals.[26]
- (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 exocytosis in noncalcifying haptophyte algae.[26]
![](http://upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Benefits_of_calcification_in_coccolithophores.jpg/500px-Benefits_of_calcification_in_coccolithophores.jpg)
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) 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.[26]
The degree by which calcification can adapt to
Silicate- or cellulose-armored functional groups such as
![](http://upload.wikimedia.org/wikipedia/commons/7/74/Energetic_effort_for_armor_construction_in_shell-forming_phytoplankton.jpg)
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
Defence against predation
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
Importance in global climate change
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Impact on the carbon cycle
Coccolithophores have both long and short term effects on the carbon cycle. The production of coccoliths requires the uptake of dissolved inorganic carbon and calcium. Calcium carbonate and carbon dioxide are produced from calcium and bicarbonate by the following chemical reaction:[99]
- Ca2+ + 2HCO−3 ⇌ CaCO3 + CO2 + H2O
Because coccolithophores are photosynthetic organisms, they are able to use some of the CO2 released in the calcification reaction for photosynthesis.[100]
However, the production of calcium carbonate drives surface alkalinity down, and in conditions of low alkalinity the CO2 is instead released back into the atmosphere.[101] As a result of this, researchers have postulated that large blooms of coccolithophores may contribute to global warming in the short term.[102] A more widely accepted idea, however, is that over the long term coccolithophores contribute to an overall decrease in atmospheric CO2 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, mediating the effects of greenhouse gas emissions.[102]
Evolutionary responses to ocean acidification
Research also suggests that
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Gephyrocapsa oceanica (scale bar is 1 μm)
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Discosphaera tubifera
Impact on microfossil record
Coccolith fossils are prominent and valuable
Of particular interest are fossils dating back to the
Impact on the oceans
The coccolithophorids help in regulating the temperature of the oceans. They thrive in warm seas and release
See also
- CLAW hypothesis
- Dimethyl sulfide
- Dimethylsulfoniopropionate
- Emiliania huxleyi virus 86
- Pleurochrysis carterae
References
- ^ ISSN 2296-7745..
Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
- ISBN 978-3-642-06016-8.
- ^ S2CID 233976784..
Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
- ^ PMID 22615387
- ^
- ^ a b 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", Transactions of the Gulf Coast Association of Geological Societies, 17: 428–480.
- ^ "Biogeography and dispersal of micro-organisms: a review emphasizing protists", Acta Protozoologica, 45 (2): 111–136, 2005
- S2CID 16601834
- ^ a b "Life at the Edge of Sight — Scott Chimileski, Roberto Kolter | Harvard University Press". www.hup.harvard.edu. Retrieved 2018-01-26.
- .
- ^ "International Nanoplankton Association".
- .
- ISBN 978-0-12-383876-6. Retrieved 30 January 2015.
- ISBN 978-3-642-06016-8.
- ISBN 978-3-642-06016-8.
- .
- .
- S2CID 34159028.
- .
- hdl:2268/246251.
- ISBN 978-3-642-06016-8.
- ^ Young, J. R. (1987). Possible Functional Interpretations of Coccolith Morphology. New York: Springer-Verlag, 305–313.
- ^ Young, J. R. (1994). "Functions of coccoliths", in Coccolithophores, eds A. Winter and W. G. Siesser (Cambridge: Cambridge University Press), 63–82.
- hdl:10453/114799.
- ^ PMID 27453937.
- .
- ^ ISBN 9780123705181.
- S2CID 21017882.
- PMID 30043404.
- .
- S2CID 22995996.
- S2CID 135347218.
- ^ Young, J. R. (1994) "Functions of coccoliths". In: Coccolithophores, Eds A. Winter and W. G. Siesser (Cambridge: Cambridge University Press), 63–82.
- S2CID 36526359.
- .
- S2CID 6227548.
- ISSN 0377-8398.
- ^
- ^ ISBN 9783540219286..
- ^ ISBN 978-0470016176
- ^ Dove, P.M.; Yoreo, J.J.; Weiner, S. (eds.). Reviews in Mineralogy and Geochemistry. Washington, D.C.: Mineralogical Society of America. pp. 189–216.
- ^ PMID 23134731
- ^
- ^ a b c Hogan, M.C. ""Coccolithophores"". In Cleveland, Cutler J. (ed.). Encyclopedia of Earth. Washington, D.C.: Environmental Information Coalition, National Council for Science and the Environment.
- ^ ISBN 9783540219286..
- ^ S2CID 9564456
- ^ a b c 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.). Origin and Evolution of Coccolithophores. Boston: Elsevier. pp. 251–285.
- ^
- ^
- doi:10.1038/s41598-019-38661-0..
Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
- ^ S2CID 27901484
- .
- doi:10.1038/ncomms10543..
Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
- doi:10.5194/bg-14-4905-2017..
Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
- .
- S2CID 4317429.
- S2CID 15482539.
- .
- S2CID 5607281.
- ^ Gitau, Beatrice (28 November 2015). "What's fueling the rise of coccolithophores in the oceans?". www.csmonitor.com. The Christian Science Monitor. Retrieved 30 November 2015.
- ^ "Viral Zone". ExPASy. Retrieved 15 June 2015.
- ^ ICTV. "Virus Taxonomy: 2014 Release". Retrieved 15 June 2015.
- ^ Largest known viral genomes Giantviruses.org. Accessed: 11 June 2020.
- ISBN 978-3-642-06016-8, archived(PDF) from the original on 2012-11-10
- ^ Morrissey, J.F.; Sumich, J.L. (2012). Introduction to the Biology of Marine Life. p. 67.
- ^
- PMID 17927770
- PMID 18824682
- (PDF) from the original on 2021-07-16.
- .
- ISBN 9780123705181.
- S2CID 84368830
- ^ "Microscopic marine plants bioengineer their environment to enhance their own growth - The Conversation". 2 August 2016.
- ^ Westbroek, P.; et al. (1983), "Calcification in Coccolithophoridae: Wasteful or Functional?", Ecological Bulletins: 291–299
- PMID 20976167
- .
- doi:10.5194/essd-10-1859-2018..
Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
- ^ PMID 21713028.
- PMID 22819465.
- PMID 23980248.
- .
- ^ .
- ^ .
- ^ doi:10.1016/j.pocean.2015.04.012..
Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
- PMID 23776586.
- .
- .
- PMID 21329207.
- Wikidata Q52718666.
- S2CID 85890446.
- .
- S2CID 135347218.
- hdl:1912/26802.
- .
- ^ hdl:1912/7739.
- ^ S2CID 90415703.
- PMID 21713029
- S2CID 85403507
- ^ PMID 14662299
- ^ S2CID 4417285
- Independent.co.uk. 22 April 2008.
- ^ "cal.mar.o". Archived from the original on 2020-12-30. Retrieved 2021-04-24.
- PMID 21713028.
- S2CID 129049029
- S2CID 4321239.
- ISBN 978-0-14-102597-1.
- S2CID 128504924.
External links
Sources of detailed information
- Nannotax3 – illustrated guide to the taxonomy of coccolithophores and other nannofossils.
- INA — International Nannoplankton Association
- Emiliania huxleyi Home Page
Introductions to coccolithophores