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Coccolith

January 31, 2023

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.

Scanning electron micrograph of Coccolithus pelagicus, plated with coccoliths

Overview

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]

  •  

    Partial cross section of a coccolithophore with coccolith layer [4]

  •  

    Coccolithophore cell surrounded by its shield of coccoliths. The coccolith-bearing cell is called the coccosphere.[5][6]

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]

Formation and composition

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]

Types

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]

 
Types of cocoliths

Shape

Coccoliths are also classified depending on shape. Common shapes include:[34][35]

  • Calyptrolith – basket-shaped with openings near the base
  • Caneolith – disc- or bowl-shaped
  • Ceratolith – horseshoe or wishbone shaped
  • Cribrilith – disc-shaped, with numerous perforations in the central area
  • Cyrtolith – convex disc shaped, may with a projecting central process
  • Discolith – ellipsoidal with a raised rim, in some cases the high rim forms a vase or cup-like structure
  • Helicolith – a placolith with a spiral margin
  • Lopadolith – basket or cup-shaped with a high rim, opening distally
  • Pentalith – pentagonal shape composed of five four-sided crystals
  • Placolith – rim composed of two plates stacked on top of one another
  • Prismatolith – polygonal, may have perforations
  • Rhabdolith – a single plate with a club-shaped central process
  • Scapholith – rhombohedral, with parallel lines in center
 
Helicoliths of Helicosphaera carteri
 
Coccosphere of Emiliania huxleyi consisting of overlapping placoliths
 
  •  

    SEM images correspond to coccolith drawings in the previous diagram
    (A) Gephyrocapsa ericsonii RCC4032 (B) Gephyrocapsa muellerae (C) Gephyrocapsa oceanica (D) Reticulofenestra parvular RCC4033; (E) Reticulofenestra parvular RCC4034; (F) Reticulofenestra parvular RCC4035; (G) Reticulofenestra parvular RCC4036; (H) Emiliania huxleyi morphotype R; (I) Emiliania huxleyi morphotype A; (J) Emiliania huxleyi morphotype B.

Function

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.

Fossil record

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. Coccoliths and related fossils are referred to as calcareous nanofossils or calcareous nannoplankton (nanoplankton).

References

  1. ^ 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", Algal Research, 1 (2): 120–133, doi:10.1016/j.algal.2012.06.002
  2. ^ Young, J. R.; Geisen, M.; Probert, I. (2005). "A review of selected aspects of coccolithophore biology with implications for paleobiodiversity estimation" (PDF). Micropaleontology. 51 (4): 267–288. doi:10.2113/gsmicropal.51.4.267.
  3. ^ a b c Haunost, Mathias; Riebesell, Ulf; D'Amore, Francesco; Kelting, Ole; Bach, Lennart T. (30 June 2021). "Influence of the Calcium Carbonate Shell of Coccolithophores on Ingestion and Growth of a Dinoflagellate Predator". Frontiers in Marine Science. Frontiers Media SA. 8. doi:10.3389/fmars.2021.664269. ISSN 2296-7745.   Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  4. ^ Irie, Takahiro; Bessho, Kazuhiro; Findlay, Helen S.; Calosi, Piero (2010-10-15). "Increasing Costs Due to Ocean Acidification Drives Phytoplankton to Be More Heavily Calcified: Optimal Growth Strategy of Coccolithophores". PLoS ONE. Public Library of Science (PLoS). 5 (10): e13436. doi:10.1371/journal.pone.0013436. ISSN 1932-6203.   Modified material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  5. ^ Aloisi, G. (6 August 2015). "Covariation of metabolic rates and cell size in coccolithophores". Biogeosciences. Copernicus GmbH. 12 (15): 4665–4692. Bibcode:2015BGeo...12.4665A. doi:10.5194/bg-12-4665-2015. ISSN 1726-4189. S2CID 6227548.
  6. ^ Henderiks, Jorijntje (2008). "Coccolithophore size rules — Reconstructing ancient cell geometry and cellular calcite quota from fossil coccoliths". Marine Micropaleontology. Elsevier BV. 67 (1–2): 143–154. Bibcode:2008MarMP..67..143H. doi:10.1016/j.marmicro.2008.01.005. ISSN 0377-8398.
  7. ^ Bown, Paul R.; Lees, Jackie A.; Young, Jeremy R. (2004). "Calcareous nannoplankton evolution and diversity through time". Coccolithophores. pp. 481–508. doi:10.1007/978-3-662-06278-4_18. ISBN 978-3-642-06016-8.
  8. ^ Hay, William W. (2004). "Carbonate fluxes and calcareous nannoplankton". Coccolithophores. pp. 509–528. doi:10.1007/978-3-662-06278-4_19. ISBN 978-3-642-06016-8.
  9. ^ 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". Deep Sea Research Part II: Topical Studies in Oceanography. 54 (5–7): 538–557. Bibcode:2007DSRII..54..538P. doi:10.1016/j.dsr2.2006.12.003.
  10. ^ Broecker, Wallace; Clark, Elizabeth (2009). "Ratio of coccolith CaCO3to foraminifera CaCO3in late Holocene deep sea sediments". Paleoceanography. 24 (3). Bibcode:2009PalOc..24.3205B. doi:10.1029/2009PA001731.
  11. ^ Klaas, Christine; Archer, David E. (2002). "Association of sinking organic matter with various types of mineral ballast in the deep sea: Implications for the rain ratio". Global Biogeochemical Cycles. 16 (4): 1116. Bibcode:2002GBioC..16.1116K. doi:10.1029/2001GB001765. S2CID 34159028.
  12. ^ 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". Progress in Oceanography. 76 (3): 217–285. Bibcode:2008PrOce..76..217H. doi:10.1016/j.pocean.2007.11.003.
  13. ^ Frankignoulle, Michel; Canon, Christine; Gattuso, Jean-Pierre (1994). "Marine calcification as a source of carbon dioxide: Positive feedback of increasing atmospheric CO2". Limnology and Oceanography. 39 (2): 458–462. Bibcode:1994LimOc..39..458F. doi:10.4319/lo.1994.39.2.0458. hdl:2268/246251.
  14. ^ Rost, Björn; Riebesell, Ulf (2004). "Coccolithophores and the biological pump: Responses to environmental changes". Coccolithophores. pp. 99–125. doi:10.1007/978-3-662-06278-4_5. ISBN 978-3-642-06016-8.
  15. ^ Young, J. R. (1987). Possible Functional Interpretations of Coccolith Morphology. New York: Springer-Verlag, 305–313.
  16. ^ Young, J. R. (1994). "Functions of coccoliths," in Coccolithophores, eds A. Winter and W. G. Siesser (Cambridge: Cambridge University Press), 63–82.
  17. ^ Raven, JA; Crawfurd, K. (2012). "Environmental controls on coccolithophore calcification". Marine Ecology Progress Series. 470: 137–166. Bibcode:2012MEPS..470..137R. doi:10.3354/meps09993.
  18. ^ a b 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). "Why marine phytoplankton calcify". Science Advances. 2 (7): e1501822. Bibcode:2016SciA....2E1822M. doi:10.1126/sciadv.1501822. PMC 4956192. PMID 27453937.
  19. ^ Müller, Marius N. (2019). "On the Genesis and Function of Coccolithophore Calcification". Frontiers in Marine Science. 6. doi:10.3389/fmars.2019.00049.
  20. ^ Hamm, Christian; Smetacek, Victor (2007). "Armor: Why, when, and How". Evolution of Primary Producers in the Sea. pp. 311–332. doi:10.1016/B978-012370518-1/50015-1. ISBN 9780123705181.
  21. ^ Brussaard, Corina P. D. (2004). "Viral Control of Phytoplankton Populations-a Review1". The Journal of Eukaryotic Microbiology. 51 (2): 125–138. doi:10.1111/j.1550-7408.2004.tb00537.x. PMID 15134247. S2CID 21017882.
  22. ^ 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). "The mutual interplay between calcification and coccolithovirus infection". Environmental Microbiology. 21 (6): 1896–1915. doi:10.1111/1462-2920.14362. PMC 7379532. PMID 30043404.
  23. ^ Haunost, Mathias; Riebesell, Ulf; Bach, Lennart T. (2020). "The Calcium Carbonate Shell of Emiliania huxleyi Provides Limited Protection Against Viral Infection". Frontiers in Marine Science. 7. doi:10.3389/fmars.2020.530757.
  24. ^ Calbet, Albert; Landry, Michael R. (2004). "Phytoplankton growth, microzooplankton grazing, and carbon cycling in marine systems". Limnology and Oceanography. 49 (1): 51–57. Bibcode:2004LimOc..49...51C. doi:10.4319/lo.2004.49.1.0051. hdl:10261/134985. S2CID 22995996.
  25. ^ 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). "Growth and mortality of coccolithophores during spring in a temperate Shelf Sea (Celtic Sea, April 2015)". Progress in Oceanography. 177: 101928. Bibcode:2019PrOce.17701928M. doi:10.1016/j.pocean.2018.02.024. S2CID 135347218.
  26. ^ Young, J. R. (1994) "Functions of coccoliths". In: Coccolithophores, Eds A. Winter and W. G. Siesser (Cambridge: Cambridge University Press), 63–82.
  27. ^ Tillmann, Urban (2004). "Interactions between Planktonic Microalgae and Protozoan Grazers1". The Journal of Eukaryotic Microbiology. 51 (2): 156–168. doi:10.1111/j.1550-7408.2004.tb00540.x. PMID 15134250. S2CID 36526359.
  28. ^ Breckels, M. N.; Roberts, E. C.; Archer, S. D.; Malin, G.; Steinke, M. (2011). "The role of dissolved infochemicals in mediating predator-prey interactions in the heterotrophic dinoflagellate Oxyrrhis marina". Journal of Plankton Research. 33 (4): 629–639. doi:10.1093/plankt/fbq114.
  29. ^ Huxley, Thomas Henry (1858). "Appendix A". Deep Sea Soundings in the North Atlantic Ocean between Ireland and Newfoundland, made in H.M.S. Cyclops, Lieut.-Commander Joseph Dayman, in June and July 1857. London: British Admiralty. pp. 63–68 [64].
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  32. ^ Fortuño, José Manuel; Cros, Lluïsa (2002-03-30). "Atlas of Northwestern Mediterranean Coccolithophores". Scientia Marina. 66 (S1): 1–182. doi:10.3989/scimar.2002.66s11. ISSN 1886-8134.
  33. ^ Malinverno, E; Dimiza, MD; Triantaphyllou, MV; Dermitzakis, MD; Corselli, C (2008). Coccolithophores of the Eastern Mediterranean sea: A look into the marine microworld. Athens: "ION" Publishing Group. ISBN 978-960-411-660-7.
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  36. ^ Bendif, El Mahdi; Probert, Ian; Díaz-Rosas, Francisco; Thomas, Daniela; van den Engh, Ger; Young, Jeremy R.; von Dassow, Peter (2016-05-24). "Recent Reticulate Evolution in the Ecologically Dominant Lineage of Coccolithophores". Frontiers in Microbiology. Frontiers Media SA. 7. doi:10.3389/fmicb.2016.00784. ISSN 1664-302X.   Modified material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.

External links

 
Wikimedia Commons has media related to Coccolith.
  • The EHUX website - site dedicated to Emiliania huxleyi, containing essays on blooms, coccolith function, etc.
  • International Nannoplankton Association site - includes an illustrated guide to coccolith terminology and several image galleries.
  • Nannotax - illustrated guide to the taxonomy of coccolithophores and other nannofossils.

January 31, 2023
coccolith, individual, plates, scales, calcium, carbonate, formed, coccolithophores, single, celled, phytoplankton, such, emiliania, huxleyi, cover, cell, surface, arranged, form, spherical, shell, called, coccosphere, scanning, electron, micrograph, pelagicus. 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 Scanning electron micrograph of Coccolithus pelagicus plated with coccoliths Contents 1 Overview 2 Formation and composition 3 Types 3 1 Shape 4 Function 5 Fossil record 6 References 7 External linksOverview EditCoccolithophores 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 Partial cross section of a coccolithophore with coccolith layer 4 Coccolithophore cell surrounded by its shield of coccoliths The coccolith bearing cell is called the coccosphere 5 6 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 Formation and composition EditCoccoliths 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 Collapsed coccosphere of Pleurochrysis carteraeTypes EditThere 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 Types of cocoliths Shape Edit Coccoliths are also classified depending on shape Common shapes include 34 35 Calyptrolith basket shaped with openings near the base Caneolith disc or bowl shaped Ceratolith horseshoe or wishbone shaped Cribrilith disc shaped with numerous perforations in the central area Cyrtolith convex disc shaped may with a projecting central process Discolith ellipsoidal with a raised rim in some cases the high rim forms a vase or cup like structure Helicolith a placolith with a spiral margin Lopadolith basket or cup shaped with a high rim opening distally Pentalith pentagonal shape composed of five four sided crystals Placolith rim composed of two plates stacked on top of one another Prismatolith polygonal may have perforations Rhabdolith a single plate with a club shaped central process Scapholith rhombohedral with parallel lines in center Helicoliths of Helicosphaera carteri Coccosphere of Emiliania huxleyi consisting of overlapping placoliths Coccolith structures of representative Noelaerhabdaceae 36 Each morphospecies is associated with a SEM image in the next diagram SEM images correspond to coccolith drawings in the previous diagram A Gephyrocapsa ericsonii RCC4032 B Gephyrocapsa muellerae C Gephyrocapsa oceanica D Reticulofenestra parvular RCC4033 E Reticulofenestra parvular RCC4034 F Reticulofenestra parvular RCC4035 G Reticulofenestra parvular RCC4036 H Emiliania huxleyi morphotype R I Emiliania huxleyi morphotype A J Emiliania huxleyi morphotype B Function EditAlthough 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 Fossil record EditBecause 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 Coccoliths and related fossils are referred to as calcareous nanofossils or calcareous nannoplankton nanoplankton References Edit 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 Algal Research 1 2 120 133 doi 10 1016 j algal 2012 06 002 Young J R Geisen M Probert I 2005 A review of selected aspects of coccolithophore biology with implications for paleobiodiversity estimation PDF Micropaleontology 51 4 267 288 doi 10 2113 gsmicropal 51 4 267 a b c Haunost Mathias Riebesell Ulf D Amore Francesco Kelting Ole Bach Lennart T 30 June 2021 Influence of the Calcium Carbonate Shell of Coccolithophores on Ingestion and Growth of a Dinoflagellate Predator Frontiers in Marine Science Frontiers Media SA 8 doi 10 3389 fmars 2021 664269 ISSN 2296 7745 Material was copied from this source which is available under a Creative Commons Attribution 4 0 International License Irie Takahiro Bessho Kazuhiro Findlay Helen S Calosi Piero 2010 10 15 Increasing Costs Due to Ocean Acidification Drives Phytoplankton to Be More Heavily Calcified Optimal Growth Strategy of Coccolithophores PLoS ONE Public Library of Science PLoS 5 10 e13436 doi 10 1371 journal pone 0013436 ISSN 1932 6203 Modified material was copied from this source which is available under a Creative Commons Attribution 4 0 International License Aloisi G 6 August 2015 Covariation of metabolic rates and cell size in coccolithophores Biogeosciences Copernicus GmbH 12 15 4665 4692 Bibcode 2015BGeo 12 4665A doi 10 5194 bg 12 4665 2015 ISSN 1726 4189 S2CID 6227548 Henderiks Jorijntje 2008 Coccolithophore size rules Reconstructing ancient cell geometry and cellular calcite quota from fossil coccoliths Marine Micropaleontology Elsevier BV 67 1 2 143 154 Bibcode 2008MarMP 67 143H doi 10 1016 j marmicro 2008 01 005 ISSN 0377 8398 Bown Paul R Lees Jackie A Young Jeremy R 2004 Calcareous nannoplankton evolution and diversity through time Coccolithophores pp 481 508 doi 10 1007 978 3 662 06278 4 18 ISBN 978 3 642 06016 8 Hay William W 2004 Carbonate fluxes and calcareous nannoplankton Coccolithophores pp 509 528 doi 10 1007 978 3 662 06278 4 19 ISBN 978 3 642 06016 8 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 Deep Sea Research Part II Topical Studies in Oceanography 54 5 7 538 557 Bibcode 2007DSRII 54 538P doi 10 1016 j dsr2 2006 12 003 Broecker Wallace Clark Elizabeth 2009 Ratio of coccolith CaCO3to foraminifera CaCO3in late Holocene deep sea sediments Paleoceanography 24 3 Bibcode 2009PalOc 24 3205B doi 10 1029 2009PA001731 Klaas Christine Archer David E 2002 Association of sinking organic matter with various types of mineral ballast in the deep sea Implications for the rain ratio Global Biogeochemical Cycles 16 4 1116 Bibcode 2002GBioC 16 1116K doi 10 1029 2001GB001765 S2CID 34159028 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 Progress in Oceanography 76 3 217 285 Bibcode 2008PrOce 76 217H doi 10 1016 j pocean 2007 11 003 Frankignoulle Michel Canon Christine Gattuso Jean Pierre 1994 Marine calcification as a source of carbon dioxide Positive feedback of increasing atmospheric CO2 Limnology and Oceanography 39 2 458 462 Bibcode 1994LimOc 39 458F doi 10 4319 lo 1994 39 2 0458 hdl 2268 246251 Rost Bjorn Riebesell Ulf 2004 Coccolithophores and the biological pump Responses to environmental changes Coccolithophores pp 99 125 doi 10 1007 978 3 662 06278 4 5 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 Raven JA Crawfurd K 2012 Environmental controls on coccolithophore calcification Marine Ecology Progress Series 470 137 166 Bibcode 2012MEPS 470 137R doi 10 3354 meps09993 a b 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 Why marine phytoplankton calcify Science Advances 2 7 e1501822 Bibcode 2016SciA 2E1822M doi 10 1126 sciadv 1501822 PMC 4956192 PMID 27453937 Muller 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3389 fmars 2020 530757 Calbet Albert Landry Michael R 2004 Phytoplankton growth microzooplankton grazing and carbon cycling in marine systems Limnology and Oceanography 49 1 51 57 Bibcode 2004LimOc 49 51C doi 10 4319 lo 2004 49 1 0051 hdl 10261 134985 S2CID 22995996 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 Growth and mortality of coccolithophores during spring in a temperate Shelf Sea Celtic Sea April 2015 Progress in Oceanography 177 101928 Bibcode 2019PrOce 17701928M doi 10 1016 j pocean 2018 02 024 S2CID 135347218 Young J R 1994 Functions of coccoliths In Coccolithophores Eds A Winter and W G Siesser Cambridge Cambridge University Press 63 82 Tillmann Urban 2004 Interactions between Planktonic Microalgae and Protozoan Grazers1 The Journal of Eukaryotic Microbiology 51 2 156 168 doi 10 1111 j 1550 7408 2004 tb00540 x PMID 15134250 S2CID 36526359 Breckels M N Roberts E C Archer S D Malin G Steinke M 2011 The role of dissolved infochemicals in mediating predator prey interactions in the heterotrophic dinoflagellate Oxyrrhis marina Journal of Plankton Research 33 4 629 639 doi 10 1093 plankt fbq114 Huxley Thomas Henry 1858 Appendix A Deep Sea Soundings in the North Atlantic Ocean between Ireland and Newfoundland made in H M S Cyclops Lieut Commander Joseph Dayman in June and July 1857 London British Admiralty pp 63 68 64 Huxley Thomas Henry 1868 On some organisms living at great depth in the North Atlantic Ocean Quarterly Journal of Microscopical Science New series 8 203 212 Sorby Henry Clifton 1861 On the organic origin of the so called Crystalloids of the chalk Annals and Magazine of Natural History Ser 3 8 45 193 200 doi 10 1080 00222936108697404 Fortuno Jose Manuel Cros Lluisa 2002 03 30 Atlas of Northwestern Mediterranean Coccolithophores Scientia Marina 66 S1 1 182 doi 10 3989 scimar 2002 66s11 ISSN 1886 8134 Malinverno E Dimiza MD Triantaphyllou MV Dermitzakis MD Corselli C 2008 Coccolithophores of the Eastern Mediterranean sea A look into the marine microworld Athens ION Publishing Group ISBN 978 960 411 660 7 Amos Winter William G Siesser 2006 Coccolithophores Cambridge University Press pp 54 58 ISBN 978 0 521 03169 1 Carmelo R Tomas 2012 Marine Phytoplankton A Guide to Naked Flagellates and Coccolithophorids Academic Press pp 161 165 ISBN 978 0 323 13827 7 Bendif El Mahdi Probert Ian Diaz Rosas Francisco Thomas Daniela van den Engh Ger Young Jeremy R von Dassow Peter 2016 05 24 Recent Reticulate Evolution in the Ecologically Dominant Lineage of Coccolithophores Frontiers in Microbiology Frontiers Media SA 7 doi 10 3389 fmicb 2016 00784 ISSN 1664 302X Modified material was copied from this source which is available under a Creative Commons Attribution 4 0 International License External links Edit Wikimedia Commons has media related to Coccolith The EHUX website site dedicated to Emiliania huxleyi containing essays on blooms coccolith function etc International Nannoplankton Association site includes an illustrated guide to coccolith terminology and several image galleries Nannotax illustrated guide to the taxonomy of coccolithophores and other nannofossils Cocco Express Coccolithophorids Expressed Sequence Tags EST amp Microarray Database Possible functions of Coccoliths Retrieved from https en wikipedia org w index php title Coccolith amp oldid 1136389357, wikipedia, wiki, book, books, library,

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