Gephyrocapsa huxleyi

{{short description|Unicellular algae responsible for the formation of chalk}}

{{more footnotes|date=April 2012}}

{{Use dmy dates|date=June 2024}}

{{Speciesbox

| image = Emiliania huxleyi.jpg

| image_alt =

| image_caption = A scanning electron micrograph of a single G. huxleyi cell

| status =

| status_system =

| status_ref = {{citation needed|date=}}

| genus = Gephyrocapsa

| species = huxleyi

| authority = (Lohmann) P.Reinhardt, 1972Reinhardt, P. (1972). Coccolithen. Kalkiges Plankton seit Jahrmillionen. Die neue Brehm Bücheri Vol. 453. pp. 1–99, 188 figures. Wittenberg Lutherstadt: A. Ziemsen Verlag

| synonyms =

| synonyms_ref =M.D. Guiry in Guiry, M.D. & Guiry, G.M. 15 November 2023. AlgaeBase. World-wide electronic publication, National University of Ireland, Galway. https://www.algaebase.org/search/species/detail/?species_id=70636 ; searched on 16 June 2024Guiry, M.D. & Guiry, G.M. (2024). AlgaeBase. World-wide electronic publication, National University of Ireland, Galway (taxonomic information republished from AlgaeBase with permission of M.D. Guiry). Gephyrocapsa huxleyi (Lohmann) P.Reinhardt, 1972. Accessed through: World Register of Marine Species at: https://www.marinespecies.org/aphia.php?p=taxdetails&id=236056 {{Webarchive|url=https://web.archive.org/web/20230804090452/https://www.marinespecies.org/aphia.php?p=taxdetails&id=236056 |date=4 August 2023 }} on 16 June 2024

}}

File:Hardangerfjord bloom 2020.jpg, Norway, May 2020]]

Gephyrocapsa huxleyi, formerly called Emiliania huxleyi, is the most abundant species of coccolithophore in modern oceans found in almost all ecosystems from the equator to sub-polar regions, and from nutrient rich upwelling zones to nutrient poor oligotrophic waters.{{Cite journal|title=The distribution of oceanic coccolithophorids in the Pacific |language=en|doi=10.1016/0011-7471(73)90059-4|volume=20|issue=4 |journal=Deep Sea Research and Oceanographic Abstracts|pages=355–374 | last1 = Okada | first1 = Hisatake |year=1973|bibcode=1973DSRA...20..355O}}Charalampopoulou, Anastasia (2011) Coccolithophores in high latitude and Polar regions: Relationships between community composition, calcification and environmental factors University of Southampton, School of Ocean and Earth Science, Doctoral Thesis, 139pp.{{Cite journal|title=Modern coccolithophoridae of the atlantic ocean—I. Placoliths and cyrtoliths |language=en|doi=10.1016/0011-7471(67)90065-4|volume=14|issue=5 |journal=Deep Sea Research and Oceanographic Abstracts|pages=561–597 | last1 = McIntyre | first1 = Andrew |year=1967|bibcode=1967DSRA...14..561M}}{{Cite journal|last1=Boeckel|first1=Babette|last2=Baumann|first2=Karl-Heinz|date=1 May 2008|title=Vertical and lateral variations in coccolithophore community structure across the subtropical frontal zone in the South Atlantic Ocean|journal=Marine Micropaleontology|volume=67|issue=3–4|pages=255–273|doi=10.1016/j.marmicro.2008.01.014|bibcode=2008MarMP..67..255B}} It is one of thousands of different photosynthetic plankton that freely drift in the photic zone of the ocean, forming the basis of virtually all marine food webs. It is studied for the extensive blooms it forms in nutrient-depleted waters after the reformation of the summer thermocline. Like other coccolithophores, G. huxleyi is a single-celled phytoplankton covered with uniquely ornamented calcite disks called coccoliths. Individual coccoliths are abundant in marine sediments although complete coccospheres are more unusual. In the case of G. huxleyi, not only the shell, but also the soft part of the organism may be recorded in sediments. It produces a group of chemical compounds that are very resistant to decomposition. These chemical compounds, known as alkenones, can be found in marine sediments long after other soft parts of the organisms have decomposed. Alkenones are most commonly used by earth scientists as a means to estimate past sea surface temperatures.

Basic facts

Emiliania huxleyi was named after Thomas Huxley and Cesare Emiliani, who were the first to examine sea-bottom sediment and discover the coccoliths within it. It is believed to have evolved approximately 270,000 years ago from the older genus Gephyrocapsa Kampter{{Cite journal|last1=Thierstein|first1=H. R.|last2=Geitzenauer|first2=K. R.|last3=Molfino|first3=B.|last4=Shackleton|first4=N. J.|date=1 July 1977|title=Global synchroneity of late Quaternary coccolith datum levels Validation by oxygen isotopes|journal=Geology|language=en|volume=5|issue=7|pages=400–404|doi=10.1130/0091-7613(1977)5<400:gsolqc>2.0.co;2|bibcode=1977Geo.....5..400T|issn=0091-7613}}{{Cite journal|last=Paasche|first=E.|title=A review of the coccolithophorid Emiliania huxleyi (Prymnesiophyceae), with particular reference to growth, coccolith formation, and calcification-photosynthesis interactions|journal=Phycologia|volume=40|issue=6|pages=503–529|doi=10.2216/i0031-8884-40-6-503.1|year=2001|bibcode=2001Phyco..40..503P |s2cid=84921998}} and became dominant in planktonic assemblages, and thus in the fossil record, approximately 70,000 years ago.{{cite journal | last1 = Bijma | first1 = J. | display-authors = etal | year = 2001 | title = Primary signal: Ecological and environmental factors—Report from Working Group 2 | doi = 10.1029/2000gc000051 | journal = Geochemistry, Geophysics, Geosystems | volume = 2 | issue = 1 | pages = n/a | bibcode = 2001GGG.....2.1003B | url = https://epic.awi.de/id/eprint/6091/1/Bij2001a.pdf | doi-access = free | access-date = 25 September 2019 | archive-date = 28 November 2020 | archive-url = https://web.archive.org/web/20201128173432/https://epic.awi.de/id/eprint/6091/1/Bij2001a.pdf | url-status = live }} The species is divided into seven morphological forms called morphotypes based on differences in coccolith structure{{Cite journal|last1=Findlay|first1=C. S|last2=Giraudeau|first2=J|date=1 December 2000|title=Extant calcareous nannoplankton in the Australian Sector of the Southern Ocean (austral summers 1994 and 1995)|journal=Marine Micropaleontology|volume=40|issue=4|pages=417–439|doi=10.1016/S0377-8398(00)00046-3|bibcode=2000MarMP..40..417F}}{{cite journal | last1 = Cook | first1 = S.S. | display-authors = etal | year = 2011 | title = Photosynthetic pigment and genetic differences between two Southern Ocean morphotypes of Emiliania Huxleyi (Haptophyta) | journal = Journal of Phycology | volume = 47 | issue = 3| pages = 615–626 | doi=10.1111/j.1529-8817.2011.00992.x| pmid = 27021991 | s2cid = 25399383 }}{{Cite journal |last1=Hagino |first1=Kyoko |last2=Bendif |first2=El Mahdi |last3=Young |first3=Jeremy R. |last4=Kogame |first4=Kazuhiro |last5=Probert |first5=Ian |last6=Takano |first6=Yoshihito |last7=Horiguchi |first7=Takeo |last8=de Vargas |first8=Colomban |last9=Okada |first9=Hisatake |date=2011-10-13 |title=NEW EVIDENCE FOR MORPHOLOGICAL AND GENETIC VARIATION IN THE COSMOPOLITAN COCCOLITHOPHORE EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE) FROM THE COX1b - ATP4 GENES 1 |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1529-8817.2011.01053.x |journal=Journal of Phycology |language=en |volume=47 |issue=5 |pages=1164–1176 |doi=10.1111/j.1529-8817.2011.01053.x |pmid=27020197 |issn=0022-3646}} (See [http://www.mikrotax.org/Nannotax3/index.php?dir=Coccolithophores/Isochrysidales/Noelaerhabdaceae/Emiliania/Emiliania%20huxleyi Nannotax] for more detail on these forms). Its coccoliths are transparent and commonly colourless, but are formed of calcite which refracts light very efficiently in the water column. This, and the high concentrations caused by continual shedding of their coccoliths makes G. huxleyi blooms easily visible from space. Satellite images show that blooms can cover areas of more than 10,000 km^2, with complementary shipboard measurements indicating that G. huxleyi is by far the dominant phytoplankton species under these conditions.{{cite journal | last1 = Holligan | first1 = P. M. | display-authors = etal | year = 1993 | title = A biogeochemical study of the coccolithophore, Emiliania huxleyi, in the North Atlantic | journal = Global Biogeochem. Cycles | volume = 7 | issue = 4| pages = 879–900 | doi = 10.1029/93GB01731 | bibcode=1993GBioC...7..879H}} This species has been an inspiration for James Lovelock's Gaia hypothesis which claims that living organisms collectively self-regulate biogeochemistry and climate at nonrandom metastable states.{{citation needed|date=March 2024}}

Abundance and distribution

Emiliania huxleyi is considered a ubiquitous species. It exhibits one of the largest temperature ranges (1–30 °C) of any coccolithophores species. It has been observed under a range of nutrient levels from oligotrophic (subtropical gyres) to eutrophic waters (upwelling zones/ Norwegian fjords).Winter, A., Jordan, R.W. & Roth, P.H., 1994. Biogeography of living coccolithophores in ocean waters. In Coccolithophores. Cambridge, United Kingdom: Cambridge University Press, pp. 161–177.{{Cite journal|last1=Hagino|first1=Kyoko|last2=Okada|first2=Hisatake|date=30 January 2006|title=Intra- and infra-specific morphological variation in selected coccolithophore species in the equatorial and subequatorial Pacific Ocean|journal=Marine Micropaleontology|volume=58|issue=3|pages=184–206|doi=10.1016/j.marmicro.2005.11.001|bibcode=2006MarMP..58..184H|hdl=2115/5820|url=https://eprints.lib.hokudai.ac.jp/dspace/bitstream/2115/5820/1/MB58-3.pdf|hdl-access=free|access-date=23 September 2019|archive-date=23 September 2019|archive-url=https://web.archive.org/web/20190923110624/https://eprints.lib.hokudai.ac.jp/dspace/bitstream/2115/5820/1/MB58-3.pdf|url-status=live}}{{Cite journal|last1=Henderiks|first1=J|last2=Winter|first2=A|last3=Elbrächter|first3=M|last4=Feistel|first4=R|last5=Plas|first5=Av der|last6=Nausch|first6=G|last7=Barlow|first7=R|date=23 February 2012|title=Environmental controls on Emiliania huxleyi morphotypes in the Benguela coastal upwelling system (SE Atlantic)|journal=Marine Ecology Progress Series|language=en|volume=448|pages=51–66|doi=10.3354/meps09535|issn=0171-8630|bibcode=2012MEPS..448...51H|doi-access=free}} Its presence in plankton communities from the surface to 200m depth indicates a high tolerance for both fluctuating and low light conditions.{{Cite journal|last1=Mohan|first1=Rahul|last2=Mergulhao|first2=Lina P.|last3=Guptha|first3=M. V. S.|last4=Rajakumar|first4=A.|last5=Thamban|first5=M.|last6=AnilKumar|first6=N.|last7=Sudhakar|first7=M.|last8=Ravindra|first8=Rasik|date=1 April 2008|title=Ecology of coccolithophores in the Indian sector of the Southern Ocean|journal=Marine Micropaleontology|volume=67|issue=1–2|pages=30–45|doi=10.1016/j.marmicro.2007.08.005|bibcode=2008MarMP..67...30M}} This extremely wide tolerance of environmental conditions is believed to be explained by the existence of a range of environmentally adapted ecotypes within the species. As a result of these tolerances its distribution ranges from the sub-Arctic to the sub-Antarctic and from coastal to oceanic habitats.Hasle, G.R., 1969. An analysis of the phytoplankton of the Pacific Southern Ocean: Abundance, composition, and distribution during the Brategg Expedition, 1947–1948, Universitetsforlaget. Within this range it is present in nearly all euphotic zone water samples and accounts for 20–50% or more of the total coccolithophore community.{{Cite journal|last1=Beaufort|first1=L.|last2=Couapel|first2=M.|last3=Buchet|first3=N.|last4=Claustre|first4=H.|last5=Goyet|first5=C.|date=4 August 2008|title=Calcite production by coccolithophores in the south east Pacific Ocean|journal=Biogeosciences|volume=5|issue=4|pages=1101–1117|doi=10.5194/bg-5-1101-2008|bibcode=2008BGeo....5.1101B|issn=1726-4189|doi-access=free}}{{cite journal | last1 = Poulton | first1 = A.J. | display-authors = etal | year = 2010 | title = Coccolithophore dynamics in non-bloom conditions during late summer in the central Iceland Basin (July–August 2007) | url = http://nora.nerc.ac.uk/id/eprint/261281/1/Poulton_L%26O_Post-print_2010.pdf | journal = Limnology and Oceanography | volume = 55 | issue = 4 | pages = 1601–1613 | doi = 10.4319/lo.2010.55.4.1601 | bibcode = 2010LimOc..55.1601P | s2cid = 53312384 | access-date = 1 July 2019 | archive-date = 11 April 2021 | archive-url = https://web.archive.org/web/20210411030555/http://nora.nerc.ac.uk/id/eprint/261281/1/Poulton_L%26O_Post-print_2010.pdf | url-status = live }}

During massive blooms (which can cover over 100,000 square kilometers), G. huxleyi cell concentrations can outnumber those of all other species in the region combined, accounting for 75% or more of the total number of photosynthetic plankton in the area. G. huxleyi blooms regionally act as an important source of calcium carbonate and dimethyl sulfide, the massive production of which can have a significant impact not only on the properties of the surface mixed layer, but also on global climate.{{Cite journal|title=A model system approach to biological climate forcing. The example of Emiliania huxleyi |language=en|doi=10.1016/0921-8181(93)90061-R|volume=8|issue=1–2 |journal=Global and Planetary Change|pages=27–46 | last1 = Westbroek | first1 = Peter |year=1993|bibcode=1993GPC.....8...27W}} The blooms can be identified through satellite imagery because of the large amount of light back-scattered from the water column, which provides a method to assess their biogeochemical importance on both basin and global scales. These blooms are prevalent in the Norwegian fjords, causing satellites to pick up "white waters", which describes the reflectance of the blooms picked up by satellites. This is due to the mass of coccoliths reflecting the incoming sunlight back out of the water, allowing the extent of G. huxleyi blooms to be distinguished in fine detail.

File:Coccolithophore samples from the Maldives.png

Extensive G. huxleyi blooms can have a visible impact on sea albedo. While multiple scattering can increase light path per unit depth, increasing absorption and solar heating of the water column, G. huxleyi has inspired proposals for geomimesis,{{cite journal | last1 = Seitz | first1 = R | year = 2011 | title = Bright water: Hydrosols, water conservation, and climate change | journal = Climatic Change | volume = 105 | issue = 3–4| pages = 365–381 | doi=10.1007/s10584-010-9965-8| arxiv = 1010.5823 | bibcode = 2011ClCh..105..365S | s2cid = 16243560 }} because micron-sized air bubbles are specular reflectors, and so in contrast to G. huxleyi, tend to lower the temperature of the upper water column. As with self-shading within water-whitening coccolithophore plankton blooms, this may reduce photosynthetic productivity by altering the geometry of the euphotic zone. Both experiments and modeling are needed to quantify the potential biological impact of such effects, and the corollary potential of reflective blooms of other organisms to increase or reduce evaporation and methane evolution by altering fresh water temperatures.

Climate change

Coccolithophores such as G. huxleyi are important to the process of ocean calcification globally; these organisms additionally play a role in long-term carbon fluxes. They fix carbon through photosynthesis, a process known as the biological carbon pump, and release carbon dioxide when coccoliths are formed. These processes therefore influence the biogeochemistry of the surface ocean.{{Cite journal |last1=Taylor |first1=Alison R. |last2=Brownlee |first2=Colin |last3=Wheeler |first3=Glen |date=2017-01-03 |title=Coccolithophore Cell Biology: Chalking Up Progress |url=https://www.annualreviews.org/content/journals/10.1146/annurev-marine-122414-034032 |journal=Annual Review of Marine Science |language=en |volume=9 |issue= |pages=283–310 |doi=10.1146/annurev-marine-122414-034032 |pmid=27814031 |bibcode=2017ARMS....9..283T |issn=1941-1405}}

Consequently, the now increasing phenomenon of ocean acidification can affect these organisms and their biological activity. As ocean acidification increases with increasing concentrations of carbon dioxide in the atmosphere, calcification by coccolithophores such as G. huxleyi can be impacted. Various studies found that increased atmospheric carbon dioxide concentrations may lead to decreased calcification by G. huxleyi. However, there is still uncertainty present regarding the degree to which calcification may be impacted, as some strains appear to be less sensitive to ocean acidification.{{Cite journal |last1=Meyer |first1=J. |last2=Riebesell |first2=U. |date=2015-03-16 |title=Reviews and Syntheses: Responses of coccolithophores to ocean acidification: a meta-analysis |url=https://bg.copernicus.org/articles/12/1671/2015/ |journal=Biogeosciences |language=English |volume=12 |issue=6 |pages=1671–1682 |doi=10.5194/bg-12-1671-2015 |doi-access=free |bibcode=2015BGeo...12.1671M |issn=1726-4170}}

As with all phytoplankton, primary production of G. huxleyi through photosynthesis is a sink of carbon dioxide. However, the production of coccoliths through calcification is a source of CO2. This means that coccolithophores, including G. huxleyi, have the potential to act as a net source of CO2 out of the ocean. Whether they are a net source or sink and how they will react to ocean acidification is not yet well understood.{{Cite journal |last1=Meyer |first1=J. |last2=Riebesell |first2=U. |date=2015-03-16 |title=Reviews and Syntheses: Responses of coccolithophores to ocean acidification: a meta-analysis |url=https://bg.copernicus.org/articles/12/1671/2015/ |journal=Biogeosciences |language=en |volume=12 |issue=6 |pages=1671–1682 |doi=10.5194/bg-12-1671-2015 |issn=1726-4189 |doi-access=free|bibcode=2015BGeo...12.1671M }}

Beyond the direct impacts to the organisms themselves, increased atmospheric carbon dioxide concentrations and therefore lower ocean pH may affect carbon cycling in the ocean. A study analyzing effects of high carbon dioxide and low pH on G. huxleyi found that PIC (particulate inorganic carbon) to POC (particulate organic carbon) ratios decreases for G. huxleyi. Under more acidic oceanic conditions, this decrease has the potential to affect export of carbon. The reasons for this decrease include a reduced calcite saturation state and disruptions to homeostasis in lower pH conditions.{{Cite journal |last1=Chauhan |first1=Nishant |last2=Rickaby |first2=Rosalind E. M. |date=2025 |title=Differential impacts of pH on growth, physiology, and elemental stoichiometry across three coccolithophore species |url=https://aslopubs.onlinelibrary.wiley.com/doi/epdf/10.1002/lno.12738 |journal=Limnology and Oceanography |language=en |volume=70 |issue=1 |pages=68–83 |doi=10.1002/lno.12738 |bibcode=2025LimOc..70...68C |issn=1939-5590|doi-access=free }}

Biogeochemical impacts

{{Unreferenced section|date=March 2024}}

=Ocean heat retention=

Scattering stimulated by G. huxleyi blooms not only causes more heat and light to be pushed back up into the atmosphere than usual, but also cause more of the remaining heat to be trapped closer to the ocean surface. This is problematic because it is the surface water that exchanges heat with the atmosphere, and G. huxleyi blooms may tend to make the overall temperature of the water column dramatically cooler over longer time periods. However, the importance of this effect, whether positive or negative, is currently being researched and has not yet been established.

Gallery

Image:Cwall99 lg.jpg|Landsat image of a 1999 G. huxleyi bloom in the English Channel off Cornwall.

Image:Bloom in the Barents Sea.jpg|G. huxleyi bloom in the Barents Sea.

See also

Notes

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