Marine primary production

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different environmental drivers on the elemental composition of phytoplankton, a comprehensive quantitative assessment of the processes is still lacking. Here, the responses of P:C and N:C ratios of marine phytoplankton have been synthesized to five major drivers (inorganic phosphorus, inorganic nitrogen, inorganic iron, irradiance, and temperature) by a meta-analysis of experimental data across 366 experiments from 104 journal articles. These results show that the response of these ratios to changes in macronutrients is consistent across all the studies, where the increase in nutrient availability is positively related to changes in P:C and N:C ratios. The results show that eukaryotic phytoplankton are more sensitive to the changes in macronutrients compared to prokaryotes, possibly due to their larger cell size and their abilities to regulate their gene expression patterns quickly. The effect of irradiance was significant and constant across all studies, where an increase in irradiance decreased both P:C and N:C. The P:C ratio decreased significantly with warming, but the response to temperature changes was mixed depending on the culture growth mode and the growth phase at the time of harvest. Along with other oceanographic conditions of the subtropical gyres (e.g., low macronutrient availability), the elevated temperature may explain why P:C is consistently low in subtropical oceans. Iron addition did not systematically change either P:C or N:C.

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vary by up to a factor of 20 between nutrient-replete and nutrient-limited cells. These studies have also shown that the C:N ratio can be modestly plastic due to nutrient limitation. A meta-analysis study by Hillebrand et al. in 2013 highlighted the importance of growth rate in determining elemental stoichiometry and showed that both C:P and N:P ratios decrease with the increasing growth rate. In 2015, Yvon-Durocher et al. investigated the role of temperature in modulating C:N:P. Although their dataset was limited to studies conducted prior to 1996, they have shown a statistically significant relationship between C:P and temperature increase. MacIntyre et al. (2002) and Thrane et al. (2016) have shown that irradiance plays an important role in controlling optimal cellular C:N and N:P ratios. Most recently, Moreno and Martiny (2018) provided a comprehensive summary of how environmental conditions regulate cellular stoichiometry from a physiological perspective.

3696: 3758: 8239: 3043: 1790: 2838: 2697: 8935: 2650: 2809: 3184: 2665: 3167: 3718:. These changes are likely to have profound effects on the physiology of phytoplankton, and observations show that competitive phytoplankton species can acclimate and adapt to changes in temperature, irradiance, and nutrients on decadal timescales. Numerous laboratory and field experiments have been conducted that study the relationship between the C:N:P ratio of phytoplankton and environmental drivers. It is, however, challenging to synthesize those studies and generalize the response of phytoplankton C:N:P to changes in environmental drivers. Individual studies employ different sets of 3714:, where the amount of inorganic nutrients, light, and temperature vary spatially and temporally. Laboratory studies show that these fluctuations trigger responses at the cellular level, whereby cells modify resource allocation in order to adapt optimally to their ambient environment. For example, phytoplankton may alter resource allocation between the P-rich biosynthetic apparatus, N-rich light-harvesting apparatus, and C-rich energy storage reserves. Under a typical future warming scenario, the global ocean is expected to undergo changes in nutrient availability, temperature, and 3785: 3203: 33: 1397: 2677: 2582: 3576: 1951: 1602: 1532: 3020: 3152: 1844: 3496: 3342: 1923: 3136: 1360: 450: 3001: 1623: 3599: 1871: 1323:

which ultimately became chloroplasts in algae and later in plants. However, while chloroplasts of the higher plants, glaucophytes, green and red algae are thought to be the result of the plastid (primary) endosymbiosis, all other groups of algae are assumed to have arisen due to the algal (secondary and tertiary) endosymbiosis (not shown), in which one eukaryotic alga was incorporated into another eukaryote. Only some branches of bacteria, eukarya, and archaea are displayed.

2567: 3483: 3309:, drought, drastic temperature shifts, etc.). They succeeded because they had the right set of traits—a mix of adaptations that were selected for in their hydro-terrestrial algal ancestors, exaptations, and the potential for co-option of a fortuitous set of genes and pathways. During the course of evolution, some members of the populations of the earliest land plants gained traits that are adaptive in terrestrial environments (such as some form of water conductance, 8961: 2434: 1027: 8949: 3227: 1635: 366: 2821: 1317: 3506: 2737: 2720: 2626: 317: 1757: 8567: 3114: 1438: 7748: 6721: 6180: 5988: 5421: 4800: 224: 2864: 3875: 3562: 369: 368: 373: 372: 367: 374: 3744:

is a powerful statistical framework for synthesizing and integrating research results obtained from independent studies and for uncovering general trends. The seminal synthesis by Geider and La Roche in 2002, as well as the more recent work by Persson et al. in 2010, has shown that C:P and N:P could

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The first eukaryote may have originated from an ancestral prokaryote that had undergone membrane proliferation, compartmentalization of cellular function (into a nucleus, lysosomes, and an endoplasmic reticulum), and the establishment of endosymbiotic relationships with an aerobic prokaryote and, in

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and denitrification. Some of these processes take place in deep water so that where there is an upwelling of cold waters, and also near estuaries where land-sourced nutrients are present, plant growth is higher. This means that the most productive areas, rich in plankton and therefore also in fish,

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The origin and evolutionary tree of life that is based on small-subunit RNA. The branches that perform oxygenic photosynthesis are labeled with 'O2'. The black arrow indicates the plastid endosymbiotic event that resulted in the origin of eukaryotic photosynthesis from cyanobacteria-like organisms,

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Research in 2019 shows these "sun-snatching bacteria" are more widespread than previously thought and could change how oceans are affected by global warming. "The findings break from the traditional interpretation of marine ecology found in textbooks, which states that nearly all sunlight in the

1279:, the building blocks for new growth, play crucial roles in regulating primary production in the ocean. Available Earth System Models suggest that ongoing ocean bio-geochemical changes could trigger reductions in ocean NPP between 3% and 10% of current values depending on the emissions scenario. 3766:

Microbial interactions outlined at the evolutionary scale, showing plant-microbe interactions occurring relatively recently compared to the more ancestral interactions among bacteria or between different microbial kingdoms. Both competitive (red) and cooperative (green) interactions within and

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The elemental stoichiometry of marine phytoplankton plays a critical role in global biogeochemical cycles through its impact on nutrient cycling, secondary production, and carbon export. Although extensive laboratory experiments have been carried out over the years to assess the influence of

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Like mangroves, seagrasses provide important nursery habitats for larval and juvenile forms of larger fish and invertebrates. The total world area of seagrass meadows is more difficult to determine than mangrove forests, but was conservatively estimated in 2003 as 177,000 square kilometres

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Primary production in the ocean can be contrasted with primary production on land. Globally the ocean and the land each produce about the same amount of primary production, but in the ocean primary production comes mainly from cyanobacteria and algae, while on land it comes mainly from

2991:. It has been estimated there are 200,000–800,000 species of which about 50,000 species have been described. Depending on the species, their sizes range from a few micrometers (μm) to a few hundred micrometers. They are specially adapted to an environment dominated by viscous forces. 3067: 3475:

Mangroves provide important nursery habitats for marine life, acting as hiding and foraging places for larval and juvenile forms of larger fish and invertebrates. Based on satellite data, the total world area of mangrove forests was estimated in 2010 as 134,257 square kilometres

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into organic carbon compounds. They are found almost everywhere on earth: in damp soil, in both freshwater and marine environments, and even on Antarctic rocks. In particular, some species occur as drifting cells floating in the ocean, and as such were amongst the first of the

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into sugars that can be used both as a source of chemical energy and of organic molecules that are used in the structural components of cells. Marine primary producers are important because they underpin almost all marine animal life by generating most of the

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and accounts for more than half the photosynthesis of the open ocean and an estimated 20% of the oxygen in the Earth's atmosphere. It is possibly the most plentiful genus on Earth: a single millilitre of surface seawater may contain 100,000 cells or more.

5312: 1890:, previously unheard of in bacteria. These proteins found in the cell membranes are capable of converting light energy to biochemical energy due to a change in configuration of the rhodopsin molecule as sunlight strikes it, causing the pumping of a 6256:

Delaux, Pierre-Marc; Hetherington, Alexander J.; Coudert, Yoan; Delwiche, Charles; Dunand, Christophe; Gould, Sven; Kenrick, Paul; Li, Fay-Wei; Philippe, Hervé; Rensing, Stefan A.; Rich, Mélanie; Strullu-Derrien, Christine; De Vries, Jan (2019).

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Béja, O.; Aravind, L.; Koonin, E.V.; Suzuki, M.T.; Hadd, A.; Nguyen, L.P.; Jovanovich, S.B.; Gates, C.M.; Feldman, R.A.; Spudich, J.L.; Spudich, E.N. (2000). "Bacterial rhodopsin: evidence for a new type of phototrophy in the sea".

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from inside out and a subsequent inflow that generates the energy. The archaeal-like rhodopsins have subsequently been found among different taxa, protists as well as in bacteria and archaea, though they are rare in complex

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that aided them in obtaining nutrients from their substrate. Furthermore, the earliest land plants had to successfully overcome a barrage of terrestrial stressors (including ultraviolet light and photosynthetically active

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De Vargas, C., Audic, S., Henry, N., Decelle, J., Mahé, F., Logares, R., Lara, E., Berney, C., Le Bescot, N., Probert, I., Carmichael, M. and 44 others (2015) "Eukaryotic plankton diversity in the sunlit ocean. Science",

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and produce half of the world's oxygen. Marine primary producers underpin almost all marine animal life by generating nearly all of the oxygen and food marine animals need to exist. Some marine primary producers are also

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traversed the world oceans collecting plankton and analysing them with contemporary molecular techniques. They found a huge range of previously unknown photosynthetic and mixotrophic algae. Among their findings were the

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Light is only able to penetrate the top 200 metres (660 ft) so this is the only part of the sea where plants can grow. The surface layers are often deficient in biologically active nitrogen compounds. The marine

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ocean is captured by chlorophyll in algae. Instead, rhodopsin-equipped bacteria function like hybrid cars, powered by organic matter when available—as most bacteria are—and by sunlight when nutrients are scarce."

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Persson, Jonas; Fink, Patrick; Goto, Akira; Hood, James M.; Jonas, Jayne; Kato, Satoshi (2010). "To be or not to be what you eat: Regulation of stoichiometric homeostasis among autotrophs and heterotrophs".

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Kim, E., Harrison, J.W., Sudek, S., Jones, M.D., Wilcox, H.M., Richards, T.A., Worden, A.Z. and Archibald, J.M.(2011) "Newly identified and diverse plastid-bearing branch on the eukaryotic tree of life".

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bacteria. In the face of global change, understanding and quantifying the mechanisms that lead to variability in C:N:P ratios are crucial in order to have an accurate projection of future climate change.

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MacIntyre, Hugh L.; Kana, Todd M.; Anning, Tracy; Geider, Richard J. (2002). "Photoacclimation of Photosynthesis Irradiance Response Curves and Photosynthetic Pigments in Microalgae and Cyanobacteria1".

1745:, when a cyanobacterium was engulfed by the eukaryote. Despite this, chloroplasts can be found in an extremely wide set of organisms, some not even directly related to each other—a consequence of many 3151: 370: 7194: 3706:

Illustration of how the five environmental drivers under a typical future climate scenario affect the cellular allocation of volume between P-rich (red), N-rich (blue), and C-rich (orange) pools.

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Flombaum, P.; Gallegos, J. L.; Gordillo, R. A.; Rincon, J.; Zabala, L. L.; Jiao, N.; Karl, D. M.; Li, W. K. W.; Lomas, M. W.; Veneziano, D.; Vera, C. S.; Vrugt, J. A.; Martiny, A. C. (2013).

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Morris, Jennifer L.; Puttick, Mark N.; Clark, James W.; Edwards, Dianne; Kenrick, Paul; Pressel, Silvia; Wellman, Charles H.; Yang, Ziheng; Schneider, Harald; Donoghue, Philip C. J. (2018).

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Gawryluk, Ryan M.R.; Del Campo, Javier; Okamoto, Noriko; Strassert, Jürgen F.H.; Lukeš, Julius; Richards, Thomas A.; Worden, Alexandra Z.; Santoro, Alyson E.; Keeling, Patrick J. (2016).

3266:—but also streptophyte algae have continuously and independently made the wet to dry transition (convergence of red and yellow). Throughout history, numerous lineages have become extinct 7468: 7028: 2797:. 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. 1886:

made a crucial discovery in the understanding of the marine carbon and energy cycles. They discovered a gene in several species of bacteria responsible for production of the protein

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as a by-product of photosynthesis induced global changes in the Earth's environment. Because oxygen was toxic to most life on Earth at the time, this led to the near-extinction of

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sequences. Many taxa, including the algal groups discussed above, are in the process of being reclassified or redefined using molecular phylogenetics. Recent developments in

5403: 3575: 7789:

Heckman, D.S., Geiser, D.M., Eidell, B.R., Stauffer, R.L., Kardos, N.L. and Hedges, S.B. (2001) "Molecular evidence for the early colonization of land by fungi and plants".

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Orth, R.J., Carruthers, T.J., Dennison, W.C., Duarte, C.M., Fourqurean, J.W., Heck, K.L., Hughes, A.R., Kendrick, G.A., Kenworthy, W.J., Olyarnik, S. and Short, F.T. (2006)

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Giri C, Ochieng E, Tieszen LL, Zhu Z, Singh A, Loveland T, et al. (2011) "Status and distribution of mangrove forests of the world using earth observation satellite data".

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International Code of Nomenclature for algae, fungi, and plants (Melbourne Code), Adopted by the Eighteenth International Botanical Congress Melbourne, Australia, July 2011

1840:. The significance of chlorophyll in converting light energy has been written about for decades, but phototrophy based on retinal pigments is just beginning to be studied. 1051: 5376:

Sparks, William B.; DasSarma, S.; Reid, I. N. (December 2006). "Evolutionary Competition Between Primitive Photosynthetic Systems: Existence of an early purple Earth?".

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Finkel, Z. V.; Beardall, J.; Flynn, K. J.; Quigg, A.; Rees, T. A. V.; Raven, J. A. (2010). "Phytoplankton in a changing world: Cell size and elemental stoichiometry".

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directly from environmental samples and avoiding the need for culturing. This has led for example, to a rapid expansion in knowledge of the abundance and diversity of

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to more complex mixed models, which makes interstudy comparisons challenging. In addition, since environmentally induced trait changes are driven by a combination of

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which surmises that original life forms on Earth were retinal-based rather than chlorophyll-based, which would have made the Earth appear purple instead of green.

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The factors limiting primary production in the ocean are also very different from those on land. The availability of water, obviously, is not an issue (though its

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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",

2676: 1531: 8393: 5117: 3166: 3019: 5977:

Faktorová, D., Dobáková, E., Peña-Diaz, P. and Lukeš, J., 2016. From simple to supercomplex: mitochondrial genomes of euglenozoan protists. F1000Research, 5.

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Gómez-Consarnau, L.; Raven, J.A.; Levine, N.M.; Cutter, L.S.; Wang, D.; Seegers, B.; Arístegui, J.; Fuhrman, J.A.; Gasol, J.M.; Sañudo-Wilhelmy, S.A. (2019).

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evolved. While the exact trait repertoire of the hypothetical last common ancestor of land plants is uncertain, it will certainly have entailed properties of

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Boyd, Philip W.; Strzepek, Robert; Fu, Feixue; Hutchins, David A. (2010). "Environmental control of open-ocean phytoplankton groups: Now and in the future".

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made up from long chains of cells, or are highly differentiated macroscopic seaweeds. They form an informal group containing about 8,000 recognized species.

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Thomas, N., Lucas, R., Bunting, P., Hardy, A., Rosenqvist, A. and Simard, M. (2017) "Distribution and drivers of global mangrove forest change, 1996–2010".

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Yvon-Durocher, Gabriel; Dossena, Matteo; Trimmer, Mark; Woodward, Guy; Allen, Andrew P. (2015). "Temperature and the biogeography of algal stoichiometry".

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The deep chlorophyll maximum (DCM) occurs at the contact where there is adequate light for photosynthesis and yet significant nutrient supply from below.

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Campbell, Neil A.; Reece, Jane B.; Urry, Lisa Andrea; Cain, Michael L.; Wasserman, Steven Alexander; Minorsky, Peter V.; Jackson, Robert Bradley (2008).

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streptophyte algae, however, did an organism whose descendants eventually conquered land on a global scale emerge: a likely branched filamentous—or even

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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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was algae, and referred to it as "blue-green algae". The more recent view is that cyanobacteria are bacteria, and hence are not even in the same

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Garcia, Catherine A.; Baer, Steven E.; Garcia, Nathan S.; Rauschenberg, Sara; Twining, Benjamin S.; Lomas, Michael W.; Martiny, Adam C. (2018).

1224:. It has been estimated that half of the world's oxygen is produced by phytoplankton. Larger autotrophs, such as the seagrasses and macroalgae ( 3695: 204:. In addition, some seagrasses, like seaweeds, can be found at depths up to 50 metres on both soft and hard bottoms of the continental shelf. 347: 4040:

Lin, I.; Liu, W. Timothy; Wu, Chun-Chieh; Wong, George T. F.; Hu, Chuanmin; Chen, Zhiqiang; Wen-Der, Liang; Yang, Yih; Liu, Kon-Kee (2003).

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Flannery, D. T.; R.M. Walter (2012). "Archean tufted microbial mats and the Great Oxidation Event: new insights into an ancient problem".

2863: 8263: 4088: 3294:(land plants) arose. In its infancy, the trajectory leading to the embryophytes was represented by the—now extinct—earliest land plants. 2696: 6980:

Martiny, Adam C.; Pham, Chau T. A.; Primeau, Francois W.; Vrugt, Jasper A.; Moore, J. Keith; Levin, Simon A.; Lomas, Michael W. (2013).

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Blankenship, R. E., Sadekar, S., and Raymond, J. (2007) "The evolutionary transition from anoxygenic to oxygenic photosynthesis". In:

2560:, they play an important role in the ecology of coral reefs. They form a (disputed) phylum containing about 7,000 recognized species. 4603: 2940:. These organisms are generally colourless and oblong in shape, typically about 20 μm long and with two flagella. Evidence from 822: 495: 8238: 7553:

Hillebrand, Helmut; Steinert, Georg; Boersma, Maarten; Malzahn, Arne; Meunier, Cédric Léo; Plum, Christoph; Ptacnik, Robert (2013).

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organisms that make their own food instead of eating other organisms. This means primary producers become the starting point in the

6030: 3757: 2820: 8413: 8198: 3734:), adaptation, and life history, stoichiometric responses of phytoplankton can be variable even amongst closely related species. 2566: 1718:

to reproduce. Their behavior is strongly influenced by environmental factors like light colour and intensity. Chloroplasts, like

1290:. They hypothesize new nutrients are flowing in from other oceans and suggest this means the Arctic Ocean may be able to support 995: 522: 6640: 7352:

Ward, B. A.; Collins, S.; Dutkiewicz, S.; Gibbs, S.; Bown, P.; Ridgwell, A.; Sauterey, B.; Wilson, J. D.; Oschlies, A. (2019).

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Wernberg, T., Krumhansl, K., Filbee-Dexter, K. and Pedersen, M.F. (2019) "Status and trends for the world’s kelp forests". In:

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seaweeds are planktonic (free-floating) and form floating drifts. Like microalgae, macroalgae (seaweeds) are technically

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is technically a protist since it is not a true plant, yet it is multicellular and can grow to 50 m (160 ft).

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cell. Chloroplasts cannot be made by the plant cell and must be inherited by each daughter cell during cell division.

1356:. These bacteria function like algae in that they can process nitrogen from the atmosphere when none is in the ocean. 8418: 7887: 7870: 7853: 7832: 6624: 5558: 4274: 3534: 3100:

covering about 25% of the world coastlines. They are among the most productive and dynamic ecosystems on Earth. Some

2808: 1874: 1209:, and a diverse group of unicellular groups. Vascular plants are also represented in the ocean by groups such as the 340: 7843: 5127: 1086:

organisms that do eat other organisms. Some marine primary producers are specialised bacteria and archaea which are

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Collins, Sinéad; Boyd, Philip W.; Doblin, Martina A. (2020). "Evolution, Microbes, and Changing Ocean Conditions".

6925:"Nutrient supply controls particulate elemental concentrations and ratios in the low latitude eastern Indian Ocean" 6341:

van der Heide, T.; van Nes, E. H.; van Katwijk, M. M.; Olff, H.; Smolders, A. J. P. (2011). Romanuk, Tamara (ed.).

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it at lower temperatures. However, the availability of light, the source of energy for photosynthesis, and mineral

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Voss, Maren; Bange, Hermann W.; Dippner, Joachim W.; Middelburg, Jack J.; Montoya, Joseph P.; Ward, Bess (2013).

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and experienced desiccation from time to time. From this "hypothetical hydro-terrestrial alga", the lineages of

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Lucking, R., Huhndorf, S., Pfister, D.H., Plata, E.R. and Lumbsch, H.T. (2009) "Fungi evolved right on track".

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and other groups. Algae encompass a diverse range of organisms, ranging from single floating cells to attached

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This visualization indicates seasonal changes in which phytoplankton types dominated over the period 1994–1998.

6330: 4817: 1789: 1548: 1539: 7227: 4476: 2719: 2321: 1750: 1519:. All chlorophylls serve as the primary means plants use to intercept light in order to fuel photosynthesis. 1258: 71:, which uses the oxidation or reduction of inorganic chemical compounds as its source of energy. Almost all 7193:

Van De Waal, Dedmer B.; Verschoor, Antonie M.; Verspagen, Jolanda MH; Van Donk, Ellen; Huisman, Jef (2010).

6487:"The marine nitrogen cycle: recent discoveries, uncertainties and the potential relevance of climate change" 5205:

Boeuf, Dominique; Audic, Stéphane; Brillet-Guéguen, Loraine; Caron, Christophe; Jeanthon, Christian (2015).

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Opacity indicates concentration of the carbon biomass. In particular, the role of the swirls and filaments (

8900: 8885: 7555:"Goldman revisited: Faster-growing phytoplankton has lower N : P and lower stoichiometric flexibility" 3710:

A key unresolved question is what determines C:N:P of individual phytoplankton. Phytoplankton grows in the

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Unlike terrestrial ecosystems, the majority of primary production in the ocean is performed by free-living

930: 925: 915: 595: 333: 293: 17: 6843:

Tyrrell, Toby (1999). "The relative influences of nitrogen and phosphorus on oceanic primary production".

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suggests diplonemids may be among the most abundant and most species-rich of all marine eukaryote groups.

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on the carbon dioxide dissolved in the water. This process uses energy from sunlight to convert water and

9025: 8895: 7901: 6607: 5634: 4822: 3835: 3830: 2050: 1165:

In a reversal of the pattern on land, in the oceans, almost all photosynthesis is performed by algae and

298: 1950: 8203: 4858: 4648:"Present and future global distributions of the marine Cyanobacteria Prochlorococcus and Synechococcus" 3676: 3262:. That said, throughout the course of evolution, algae from various other lineages have colonized land 2412: 2144: 2019: 985: 883: 589: 583: 7729:

Hassani, M.A., Durán, P. and Hacquard, S. (2018) "Microbial interactions within the plant holobiont".

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to characterize the effects of the environmental driver(s) on elemental ratios, ranging from a simple

2160: 1344: 1282:

In 2020 researchers reported that measurements over the last two decades of primary production in the

32: 8985: 8405: 3683:, with additional influences from biological diversity and secondary processing of organic matter by 3333:. What is also certain is that the last common ancestor of land plants had traits of algal ancestry. 2451: 2149: 990: 5604: 5402:

Javed, M.R., Bilal, M.J., Ashraf, M.U.F., Waqar, A., Mehmood, M.A., Saeed, M. and Nashat, N. (2019)

4788: 4126:"Biotic and Human Vulnerability to Projected Changes in Ocean Biogeochemistry over the 21st Century" 8964: 8757: 8752: 8048: 3648: 3246:

The diagram on the right shows an evolutionary scenario for the conquest of land by streptophytes.

1714:. They are highly dynamic—they circulate and are moved around within plant cells, and occasionally 965: 893: 888: 871: 716: 577: 475: 6484: 5313:"Early evolution of purple retinal pigments on Earth and implications for exoplanet biosignatures" 8990: 8665: 8476: 8370: 8098: 8088: 7683:"The impact of irradiance on optimal and cellular nitrogen to phosphorus ratios in phytoplankton" 4628: 3561: 3270:. Terrestrial algae of various taxonomic affiliations dwell on rock surfaces and form biological 2922: 2890: 2397: 2366: 2043: 1911: 1483:. It is the presence and relative abundance of chlorophyll that gives plants their green colour. 975: 490: 9005: 8491: 8481: 8183: 8168: 6314: 5654: 5599: 3088:. Seaweeds usually grow in shallow coastal waters where they are anchored to the seafloor by a 2371: 2244: 1896: 1851: 1807: 795: 571: 532: 500: 5207:"MicRhoDE: a curated database for the analysis of microbial rhodopsin diversity and evolution" 4422: 3854: 3647:(P) in exported organic matter expressed in terms of the C:N:P ratio helps determine how much 1246:, but the vast majority of free-floating production takes place within microscopic organisms. 8910: 8837: 8772: 8637: 8612: 8587: 8471: 8218: 7998: 6103: 4897:"The light-driven proton pump proteorhodopsin enhances bacterial survival during tough times" 4758: 4262: 3727: 2910: 1644:

some cases, a photosynthetic prokaryote to form mitochondria and chloroplasts, respectively.

1044: 1031: 980: 711: 238: 91: 7242:

Irwin, Andrew J.; Finkel, Zoe V.; Müller-Karger, Frank E.; Troccoli Ghinaglia, Luis (2015).

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Mann, K.H. 1973. Seaweeds: their productivity and strategy for growth. Science 182: 975-981.

5548: 5263: 4202:"Changes in phytoplankton concentration now drive increased Arctic Ocean primary production" 8645: 8607: 7938: 7694: 7647: 7611: 7566: 7527: 7483: 7425: 7365: 7310: 7255: 7206: 7132: 7089: 7080:

Moreno, Allison R.; Martiny, Adam C. (2018). "Ecological Stoichiometry of Ocean Plankton".

7043: 6993: 6982:"Strong latitudinal patterns in the elemental ratios of marine plankton and organic matter" 6936: 6852: 6786: 6747: 6692: 6354: 6270: 6210: 5942: 5815: 5773: 5700: 5591: 5447: 5385: 5334: 5171: 5075: 5021: 4962: 4659: 4579: 4499: 4053: 3820: 3719: 3652: 3314: 2918: 2902: 2893:

have allowed the evolutionary relationship of species to be established by analyzing their

2886: 2589: 2301: 2065: 1800: (2) it changes its configuration so a proton is expelled from the cell 1773: 1707: 1460: 1181:

are important photosynthetizers in both oceanic and terrestrial ecosystems, and while some

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cycles. They stabilize coastal areas and can provide habitats for marine animals. The term

1139: 907: 809: 565: 470: 268: 4328: 1802: (3) the chemical potential causes the proton to flow back to the cell 1601: 8: 8867: 8521: 8330: 8288: 8058: 8018: 7469:"Redfield revisited: Variability of C:N:P in marine microalgae and its biochemical basis" 7322: 7101: 7029:"Redfield revisited: Variability of C:N:P in marine microalgae and its biochemical basis" 6340: 5582:

Yool, A.; Tyrrell, T. (2003). "Role of diatoms in regulating the ocean's silicon cycle".

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Lee, DW (2007) Nature's palette - the science of plant color. University of Chicago Press

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are any coloured material in plant or animal cells. All biological pigments selectively

48:, while blue indicates where there are few phytoplankton. – NASA Earth Observatory 2019. 8787: 8782: 8650: 8592: 8486: 8335: 8253: 8138: 8118: 8028: 7971: 7663: 7584: 7499: 7449: 7386: 7353: 7334: 7278: 7243: 7148: 7059: 7009: 6957: 6924: 6868: 6804: 6710: 6511: 6486: 6377: 6342: 6296: 6233: 6198: 6164: 6139: 6055: 5836: 5799: 5723: 5617: 5471: 5404:"Microalgae as a Feedstock for Biofuel Production: Current Status and Future Prospects" 5358: 5324: 5239: 5206: 5099: 5034: 5009: 4985: 4950: 4923: 4896: 4682: 4647: 4533: 4515: 4303: 4237: 4152: 4125: 4069: 3422:. All of these habitats are able to sequester large quantities of carbon and support a 3330: 3209: 3089: 2984: 2882: 2636: 2390: 2326: 2208: 1452: 1390: 1386: 1272: 1233: 1151: 1135: 1103: 1091: 935: 703: 687: 682: 605: 321: 273: 253: 108: 76: 41: 37: 7354:"Considering the Role of Adaptive Evolution in Models of the Ocean and Climate System" 6681:"A meta-analysis on environmental drivers of marine phytoplankton C : N : P" 6028:

A place in the sun - Algae is the crop of the future, according to researchers in Geel

5907:

Bork, P., Bowler, C., De Vargas, C., Gorsky, G., Karsenti, E. and Wincker, P. (2015) "

5885: 5860: 4591: 4201: 3986: 3961: 1291: 9020: 9015: 8797: 8675: 8622: 8541: 8208: 8173: 8113: 8073: 8043: 8013: 7924: 7883: 7866: 7849: 7839: 7828: 7818: 7712: 7659: 7539: 7441: 7409: 7391: 7338: 7326: 7283: 7105: 6962: 6759: 6714: 6620: 6571: 6539: 6516: 6382: 6343:"Positive feedbacks in seagrass ecosystems: evidence from large-scale empirical data" 6300: 6288: 6238: 6169: 6109: 6082: 5960: 5890: 5841: 5728: 5564: 5554: 5527: 5493: 5463: 5459: 5411: 5362: 5350: 5269: 5244: 5226: 5187: 5091: 5047: 5039: 4990: 4951:"Microbial rhodopsins are major contributors to the solar energy captured in the sea" 4928: 4877: 4762: 4710: 4687: 4595: 4470: 4430: 4383: 4270: 4241: 4229: 4221: 4157: 3991: 3936: 3915: 3741: 3668: 3656: 3530: 3515: 3455: 2789: 2772: 2762: 2727: 2438: 2331: 2112: 2038: 1855: 1427: 866: 645: 480: 243: 80: 7667: 7588: 7503: 7152: 7063: 6808: 5621: 5475: 4519: 4421:

Walsh PJ, Smith S, Fleming L, Solo-Gabriele H, Gerwick WH, eds. (2 September 2011).

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Garcia-Mendoza, E. and Ocampo-Alvarez, H. (2011) "Photoprotection in the brown alga

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of microorganisms, such as the algal groups discussed above, was inferred and their

1396: 426: 8905: 8597: 8501: 8453: 8365: 8258: 7956: 7798: 7773: 7738: 7702: 7655: 7619: 7574: 7535: 7491: 7453: 7433: 7381: 7373: 7318: 7273: 7263: 7222: 7214: 7175: 7140: 7097: 7051: 7013: 7001: 6952: 6944: 6903: 6872: 6860: 6794: 6755: 6700: 6591: 6567: 6506: 6498: 6372: 6362: 6326: 6278: 6228: 6218: 6159: 6151: 6051: 6010: 5978: 5950: 5912: 5880: 5872: 5831: 5823: 5781: 5718: 5708: 5609: 5455: 5342: 5234: 5218: 5179: 5103: 5083: 5029: 4980: 4970: 4918: 4908: 4867: 4754: 4677: 4667: 4587: 4541: 4507: 4404: 4379: 4350: 4324: 4315:

Hohmann-Marriott, M.F. and Blankenship, R.E. (2011) "Evolution of photosynthesis".

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camouflaged to look like floating seaweed live in kelp forests and seagrass meadows

3581: 3538: 3287: 2794: 2703: 2533: 2033: 1992: 1340: 1119:

and food that provide other organisms with the chemical energy they need to exist.

1071: 832: 726: 721: 103: 56: 6775:"Redfield revisited: 1. Regulation of nitrate, phosphate, and oxygen in the ocean" 5982: 5183: 5087: 4042:"New evidence for enhanced ocean primary production triggered by tropical cyclone" 3495: 3341: 2971:

are the microscopic types of algae, not visible to the naked eye. They are mostly

1960: 8875: 8842: 8703: 8655: 8531: 8506: 8315: 8268: 8193: 8038: 8023: 6616: 6595: 6367: 6034: 4913: 4511: 4142: 3825: 3810: 3711: 3672: 3567: 3511: 3427: 3407: 3403: 2929: 2750: 2614: 2572: 2557: 2514: 2356: 2254: 2213: 2203: 2096: 1568: 1410: 1348: 1295: 1170: 960: 827: 760: 748: 677: 551: 411: 288: 283: 258: 197: 161: 157: 5745:"Biogeography and dispersal of micro-organisms: a review emphasizing protists", 1922: 9010: 8739: 8693: 8688: 8660: 8627: 8496: 8355: 8283: 8273: 8123: 8068: 7986: 7981: 7976: 7752: 6981: 6948: 6887: 6725: 6184: 5992: 5785: 5425: 4804: 3805: 3451: 3423: 3326: 3279: 3189: 3135: 2827: 2776: 2532:

live most of their lives as single cells or are filamentous, while others form

2518: 2316: 2296: 2264: 2169: 2101: 2011: 1997: 1727: 1692: 1664: 1575: 1464: 1359: 1190: 1111: 1107: 1099: 970: 940: 920: 861: 775: 658: 653: 514: 248: 233: 120: 112: 68: 64: 60: 7742: 7579: 7554: 7495: 7195:"Climate-driven changes in the ecological stoichiometry of aquatic ecosystems" 7144: 7055: 6283: 6258: 5955: 5930: 5827: 5346: 5222: 4298:, eds P. G. Falkowski and A. N. Knoll, New York: Academic Press, pages 21–35. 3675:

and burial. On shorter timescales, the average stoichiometry of exported bulk

3659:. On geologic timescales, the N:P ratio reflects the relative availability of 3602: 2481:

is an informal term for a widespread and diverse collection of photosynthetic

1368:. Cyanobacteria were the first organisms to release oxygen via photosynthesis. 8979: 8953: 8847: 8812: 8713: 8708: 8683: 8213: 8178: 8153: 8003: 7966: 6636: 5931:"Morphological Identification and Single-Cell Genomics of Marine Diplonemids" 5568: 5354: 5230: 5043: 4282: 4225: 3879: 3737: 3688: 3680: 3620: 3459: 3081: 3034: 3030: 3006: 2976: 2941: 2670:

They are a major algae group generating about 20% of world oxygen production.

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and several colourful pigments that absorb as much light energy as possible.

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Shevela, D. (2011) "Adventures with cyanobacteria: a personal perspective".

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are almost exclusively marine and are found in large numbers throughout the

1843: 449: 8939: 8718: 8698: 8602: 8340: 8223: 8143: 8103: 8093: 8063: 7716: 7445: 7395: 7330: 7287: 7109: 6966: 6892:"The role of phytoplankton diversity in the emergent oceanic stoichiometry" 6520: 6502: 6386: 6292: 6242: 6173: 5964: 5845: 5732: 5467: 5248: 5191: 5095: 5051: 4994: 4975: 4932: 4881: 4872: 4853: 4766: 4691: 4599: 4233: 4161: 3995: 3446:. These plants have adapted to the high salinity of the ocean environment. 3439: 3322: 3275: 3247: 3236: 2988: 2975:

species which exist as individuals or in chains or groups, though some are

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Phototrophic metabolism relies on one of three energy-converting pigments:

1719: 1696: 1679:, converts it, and stores it in the energy-storage molecules while freeing 1283: 1197:

are significant contributors to primary production in the ocean, including

1143: 753: 667: 621: 559: 441: 263: 7241: 6155: 5894: 5861:"The winds of (evolutionary) change: breathing new life into microbiology" 4745:

Grotewold, E. (2006). "The Genetics and Biochemistry of Floral Pigments".

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Hoek, Christiaan; den Hoeck, Hoeck Van; Mann, David; Jahns, H.M. (1995).

4538:"Understand the evolutionary mechanisms and environmental limits of life" 4065: 3731: 3684: 3631:) of primary production in the surface ocean plays a crucial role in the 3364: 3360: 3291: 3283: 3251: 3127: 3097: 3070: 2972: 2784: 2754: 2602: 2529: 2506: 2498: 2407: 2402: 2351: 2291: 2283: 2194: 1973: 1833: 1821: 1765: 1761: 1711: 1668: 1648: 1615: 1484: 1472: 1468: 1431: 1237: 1147: 1083: 817: 790: 785: 780: 770: 740: 626: 485: 429:

features) appear important in maintaining high biodiversity in the ocean.

215: 177: 173: 127: 102:. Together these form the principal primary producers at the base of the 7437: 6822: 1726:, which is thought to be inherited from their ancestor—a photosynthetic 8832: 8767: 8556: 8536: 8320: 8293: 8078: 7991: 3715: 3644: 3426:

range of larger and smaller animal life. Marine plants can be found in

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Most marine primary production is generated by a diverse collection of

67:, which uses light as its source of energy, but it also occurs through 8948: 7751: