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Phytoplankton
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{{short description| Autotrophic members of the plankton ecosystem }}{{Use dmy dates|date=April 2012}}
missing image!
- Diatoms through the microscope.jpg -
Diatoms are one of the most common types of phytoplankton.
Phytoplankton {{IPAc-en|ˌ|f|aɪ|t|oʊ|ˈ|p|l|æ|ŋ|k|t|ə|n}} are the autotrophic (self-feeding) components of the plankton community and a key part of oceans, seas and freshwater basin ecosystems. The name comes from the Greek words φυτόν (phyton), meaning "plant", and πλανκτός (planktos), meaning "wanderer" or "drifter".BOOK
, Thurman
, H. V.
, 2007
, Introductory Oceanography
, Academic Internet Publishers
,
, 978-1-4288-3314-2
, {{page needed|date=February 2016}} Most phytoplankton are too small to be individually seen with the unaided eye. However, when present in high enough numbers, some varieties may be noticeable as colored patches on the water surface due to the presence of chlorophyll within their cells and accessory pigments (such as phycobiliproteins or xanthophylls) in some species. About 1% of the global biomass is due to phytoplankton.JOURNAL, Bidle KD, Falkowski PG, Cell death in planktonic, photosynthetic microorganisms, Nature Reviews. Microbiology, 2, 8, 643–655, August 2004, 15263899, 10.1038/nrmicro956,

Types

Phytoplankton are extremely diverse, varying from photosynthesising bacteria (cyanobacteria), to plant-like diatoms, to armour-plated coccolithophores.Lindsey, R., Scott, M. and Simmon, R. (2010) "What are phytoplankton". NASA Earth Observatory.File:Phytoplankton types.jpg|left|thumb|720px|{{spaces|10}}cyanobacteria{{spaces|20}}diatom{{spaces|20}}dinoflagellate{{spaces|26}}green algae{{spaces|20}}(drawings not to scale)}}{{clear}}

Ecology

{{multiple image
| align = right
| direction = vertical
| width = 220
| header =
| image1 = Phytopla.jpg
| alt1 =
| caption1 = Phytoplankton come in many shapes and sizes.
| image2 = Phytoplankton - the foundation of the oceanic food chain.jpg
| alt2 =
| caption2 = Phytoplankton are the foundation of the oceanic food chain.
| image3 = Spring Bloom Colors the Pacific Near Hokkaido.jpg
| alt3 =
| caption3 = When two currents collide (here the Oyashio and Kuroshio currents) they create eddies. Phytoplankton concentrates along the boundaries of the eddies, tracing the motion of the water.
| image4 = Cwall99 lg.jpg
| alt4 =
| caption4 = Algal bloom off south west England.
}}

Carbon

Phytoplankton are photosynthesizing microscopic biotic organisms that inhabit the upper sunlit layer of almost all oceans and bodies of fresh water on Earth. They are agents for "primary production", the creation of organic compounds from carbon dioxide dissolved in the water, a process that sustains the aquatic food web.WEB, Ghosal; Rogers; Wray, S.; M.; A., The Effects of Turbulence on Phytoplankton,weblink Aerospace Technology Enterprise, NTRS, 16 June 2011, Phytoplankton obtain energy through the process of photosynthesis and must therefore live in the well-lit surface layer (termed the euphotic zone) of an ocean, sea, lake, or other body of water. Phytoplankton account for about half of all photosynthetic activity on Earth.JOURNAL, Michael J. Behrenfeld, etal, Biospheric primary production during an ENSO transition, Science, 2001-03-30, 291, 5513, 2594–7, 10.1126/science.1055071, 11283369, "NASA Satellite Detects Red Glow to Map Global Ocean Plant Health" NASA, 28 May 2009.WEB,weblink Satellite Sees Ocean Plants Increase, Coasts Greening, 9 June 2014, NASA, 2 March 2005, Their cumulative energy fixation in carbon compounds (primary production) is the basis for the vast majority of oceanic and also many freshwater food webs (chemosynthesis is a notable exception).While almost all phytoplankton species are obligate photoautotrophs, there are some that are mixotrophic and other, non-pigmented species that are actually heterotrophic (the latter are often viewed as zooplankton). Of these, the best known are dinoflagellate genera such as Noctiluca and Dinophysis, that obtain organic carbon by ingesting other organisms or detrital material.

Oxygen production

Phytoplankton absorb energy from the Sun and nutrients from the water to produce their own food. In the process of photosynthesis, phytoplankton release molecular oxygen ({{chem|O|2}}) into the water. It is estimated that between 50% and 85% of the world's oxygen is produced via phytoplankton photosynthesis.NEWS,weblink How much do oceans add to world's oxygen?, June 8, 2015, Earth & Sky, 2016-04-04, WEB,weblink Phytoplankton levels dropping (NASA), Youtube, NEWS,weblink Source of Half Earth's Oxygen Gets Little Credit, Roach, John, June 7, 2004, National Geographic News, 2016-04-04, JOURNAL, Ryther, John H., 25 July 1970, Biological Sciences: Is the World's Oxygen Supply Threatened ?,weblink Nature, en, 227, 5256, 374–375, 10.1038/227374a0, 0028-0836, JOURNAL, 10.1029/2003GL017141, 2003GeoRL..30.1718L, 30, 13, New evidence for enhanced ocean primary production triggered by tropical cyclone, 2003, Geophysical Research Letters, Lin, I., Liu, W. Timothy, Wu, Chun-Chieh, Wong, George T. F., Hu, Chuanmin, Chen, Zhiqiang, Wen-Der, Liang, Yang, Yih, Liu, Kon-Kee,weblink The rest is produced via photosynthesis on land by plants. Furthermore, phytoplankton photosynthesis has controlled the atmospheric {{chem|C|O|2}}/{{chem|O|2}} balance since the early Precambrian Eon.JOURNAL, Tappan, Helen, April 1968, Primary production, isotopes, extinctions and the atmosphere, Palaeogeography, Palaeoclimatology, Palaeoecology, 4, 3, 187–210, 1968PPP.....4..187T, 10.1016/0031-0182(68)90047-3, (See Biological pump.)

Minerals

Phytoplankton are crucially dependent on minerals. These are primarily macronutrients such as nitrate, phosphate or silicic acid, whose availability is governed by the balance between the so-called biological pump and upwelling of deep, nutrient-rich waters. Phytoplankton nutrient composition drives and is driven by the Redfield ratio of macronutrients generally available throughout the surface oceans. However, across large regions of the World Ocean such as the Southern Ocean, phytoplankton are also limited by the lack of the micronutrient iron. This has led to some scientists advocating iron fertilization as a means to counteract the accumulation of human-produced carbon dioxide (CO2) in the atmosphere.NEWS, M., Richtel, Recruiting Plankton to Fight Global Warming, New York Times, 1 May 2007,weblink Large-scale experiments have added iron (usually as salts such as iron sulphate) to the oceans to promote phytoplankton growth and draw atmospheric CO2 into the ocean. However, controversy about manipulating the ecosystem and the efficiency of iron fertilization has slowed such experiments.JOURNAL, Monastersky, Richard, Iron versus the Greenhouse: Oceanographers Cautiously Explore a Global Warming Therapy, Science News, 148, 14, 1995, 220–1, 10.2307/4018225, 4018225,

B Vitamins

Phytoplankton depend on B Vitamins for survival. Areas in the ocean have been identified as having a major lack of some B Vitamins, and correspondingly, phytoplankton.WEB,weblink Existence of vitamin 'deserts' in the ocean confirmed, ScienceDaily, Sergio, Sañudo-Wilhelmy, 2012-06-23,

Temperature

The effects of anthropogenic warming on the global population of phytoplankton is an area of active research. Changes in the vertical stratification of the water column, the rate of temperature-dependent biological reactions, and the atmospheric supply of nutrients are expected to have important effects on future phytoplankton productivity.JOURNAL, Henson, S. A., Sarmiento, J. L., Dunne, J. P., Bopp, L., Lima, I., Doney, S. C., John, J., Beaulieu, C., 2010, Detection of anthropogenic climate change in satellite records of ocean chlorophyll and productivity, Biogeosciences, 7, 2, 621–40, 10.5194/bg-7-621-2010, JOURNAL, Steinacher, M., Joos, F., Frölicher, T. L., Bopp, L., Cadule, P., Cocco, V., Doney, S. C., Gehlen, M., Lindsay, K., 2010, Projected 21st century decrease in marine productivity: a multi-model analysis, Biogeosciences, 7, 3, 979–1005, 10.5194/bg-7-979-2010, Moore, J. K., Schneider, B., Segschneider, J.,

pH

The effects of anthropogenic ocean acidification on phytoplankton growth and community structure has also received considerable attention. Phytoplankton such as coccolithophores contain calcium carbonate cell walls that are sensitive to ocean acidification. Because of their short generation times, evidence suggests some phytoplankton can adapt to changes in pH induced by increased carbon dioxide on rapid time-scales (months to years).JOURNAL, Collins, Sinéad, Rost, Björn, Rynearson, Tatiana A., 2013-11-25, Evolutionary potential of marine phytoplankton under ocean acidification, Evolutionary Applications, en, 7, 1, 140–155, 10.1111/eva.12120, 1752-4571, 3894903, 24454553, JOURNAL, Lohbeck, Kai T., Riebesell, Ulf, Reusch, Thorsten B. H., 2012-04-08, Adaptive evolution of a key phytoplankton species to ocean acidification, Nature Geoscience, En, 5, 5, 346–351, 10.1038/ngeo1441, 1752-0894,

Food web

Phytoplankton serve as the base of the aquatic food web, providing an essential ecological function for all aquatic life. Under future conditions of anthropogenic warming and ocean acidification, changes in phytoplankton mortality may be significant.{{citation needed|date=January 2019}}{{elucidate|date=January 2019}} One of the many food chains in the ocean – remarkable due to the small number of links – is that of phytoplankton sustaining krill (a crustacean similar to a tiny shrimp), which in turn sustain baleen whales.

Structural and functional diversity

The term phytoplankton encompasses all photoautotrophic microorganisms in aquatic food webs. However, unlike terrestrial communities, where most autotrophs are plants, phytoplankton are a diverse group, incorporating protistan eukaryotes and both eubacterial and archaebacterial prokaryotes. There are about 5,000 known species of marine phytoplankton.BOOK, Manual on Harmful Marine Microalgae, Hallegraeff, G.M., Unesco, 2003, 978-92-3-103871-6, Hallegraeff, Gustaaf M., 25–49, Harmful algal blooms: a global overview, Anderson, Donald Mark, Cembella, Allan D., Enevoldsen, Henrik O.,weblink How such diversity evolved despite scarce resources (restricting niche differentiation) is unclear.JOURNAL, Hutchinson, G. E., 1961, The Paradox of the Plankton, The American Naturalist, 95, 882, 137–45, 10.1086/282171, In terms of numbers, the most important groups of phytoplankton include the diatoms, cyanobacteria and dinoflagellates, although many other groups of algae are represented. One group, the coccolithophorids, is responsible (in part) for the release of significant amounts of dimethyl sulfide (DMS) into the atmosphere. DMS is oxidized to form sulfate which, in areas where ambient aerosol particle concentrations are low, can contribute to the population of cloud condensation nuclei, mostly leading to increased cloud cover and cloud albedo according to the so-called CLAW Hypothesis.JOURNAL, Charlson, Robert J., Lovelock, James E., Andreae, Meinrat O., Warren, Stephen G., 1987, Oceanic phytoplankton, atmospheric sulphur, cloud albedo and climate, Nature, 326, 6114, 655–61, 1987Natur.326..655C, 10.1038/326655a0, JOURNAL, Quinn, P. K., Bates, T. S., 2011, The case against climate regulation via oceanic phytoplankton sulphur emissions, Nature, 480, 7375, 51–6, 2011Natur.480...51Q, 10.1038/nature10580, 22129724,weblink Different types of phytoplankton support different trophic levels within varying ecosystems. In oligotrophic oceanic regions such as the Sargasso Sea or the South Pacific Gyre, phytoplankton is dominated by the small sized cells, called picoplankton and nanoplankton (also referred to as picoflagellates and nanoflagellates), mostly composed of cyanobacteria (Prochlorococcus, Synechococcus) and picoeucaryotes such as Micromonas. Within more productive ecosystems, dominated by upwelling or high terrestrial inputs, larger dinoflagellates are the more dominant phytoplankton and reflect a larger portion of the biomass.JOURNAL, Calbet, A., 2008, The trophic roles of microzooplankton in marine systems, ICES Journal of Marine Science, 65, 3, 325–31, 10.1093/icesjms/fsn013,

Growth strategy

In the early twentieth century, Alfred C. Redfield found the similarity of the phytoplankton’s elemental composition to the major dissolved nutrients in the deep ocean.BOOK, Redfield, Alfred C., 1934, On the Proportions of Organic Derivatives in Sea Water and their Relation to the Composition of Plankton, 176–92, Johnstone, James, Daniel, Richard Jellicoe, James Johnstone Memorial Volume, Liverpool, University Press of Liverpool, 13993674, Redfield proposed that the ratio of carbon to nitrogen to phosphorus (106:16:1) in the ocean was controlled by the phytoplankton’s requirements, as phytoplankton subsequently release nitrogen and phosphorus as they are remineralized. This so-called “Redfield ratio” in describing stoichiometry of phytoplankton and seawater has become a fundamental principle to understand marine ecology, biogeochemistry and phytoplankton evolution.JOURNAL, Arrigo, Kevin R., Marine microorganisms and global nutrient cycles, Nature, 437, 7057, 349–55, 2005, 16163345, 10.1038/nature04159, 2005Natur.437..349A, However, the Redfield ratio is not a universal value and it may diverge due to the changes in exogenous nutrient deliveryJOURNAL, Fanning, Kent A., Influence of atmospheric pollution on nutrient limitation in the ocean, Nature, 339, 6224, 1989, 460–63, 1989Natur.339..460F, 10.1038/339460a0, and microbial metabolisms in the ocean, such as nitrogen fixation, denitrification and anammox.The dynamic stoichiometry shown in unicellular algae reflects their capability to store nutrients in an internal pool, shift between enzymes with various nutrient requirements and alter osmolyte composition.BOOK, Robert Warner, Sterner, James J., Elser, 2002, Ecological Stoichiometry: The Biology of Elements from Molecules to the Biosphere, Princeton University Press, 978-0-691-07491-7, {{page needed|date=February 2016}}JOURNAL, Klausmeier, Christopher A., Litchman, Elena, Levin, Simon A., Phytoplankton growth and stoichiometry under multiple nutrient limitation, Limnology and Oceanography, 49, 4 Part 2, 2004, 1463–70, 10.4319/lo.2004.49.4_part_2.1463, 2004LimOc..49.1463K, Different cellular components have their own unique stoichiometry characteristics, for instance, resource (light or nutrients) acquisition machinery such as proteins and chlorophyll contain a high concentration of nitrogen but low in phosphorus. Meanwhile, growth machinery such as ribosomal RNA contains high nitrogen and phosphorus concentrations.Based on allocation of resources, phytoplankton is classified into three different growth strategies, namely survivalist, bloomerJOURNAL, Klausmeier, Christopher A., Litchman, Elena, Daufresne, Tanguy, Levin, Simon A., Optimal nitrogen-to-phosphorus stoichiometry of phytoplankton, Nature, 429, 6988, 171–4, 2004, 15141209, 2004Natur.429..171K, 10.1038/nature02454, and generalist. Survivalist phytoplankton has a high ratio of N:P (>30) and contains an abundance of resource-acquisition machinery to sustain growth under scarce resources. Bloomer phytoplankton has a low N:P ratio (

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