Knowledge (XXG)

Lipid pump

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1228: 1284: 908:. Individuals enter diapause and migrate deeper in the water column to overwinter below the thermocline. During diapause they survive on stored lipid reserves that are generated at the end of their time at the surface when nutrients are widely available. The seasonal end of diapause must be closely timed with the beginning of the spring phytoplankton bloom to enable acquisition of food to permit proper egg development and hatching. If the timing is disrupted, eggs that are hatched during diapause will have limited growth time and a lower likelihood of surviving overwintering, as thus is an example of 1300:($ 413 million). Pollock alone is the largest fishery in the US based on volume, but is also the second largest fishery in the world supporting 2–5% of the global fishery production. Not only do millions of people rely on fish for subsistence, but recreational fishing is one of the most popular activities in the US. Recreational fishing contributes about $ 202 million to the US economy. Changes in the abundance and distribution of copepods could drastically affect the economic livelihoods of millions of people connected to the fishing industry or who rely on fishing as a primary source of protein. 1473:
accurate ways to measure both mortality and respiration rates of overwintering zooplankton are being conducted in recent work, which are the two factors that primarily control the amount of lipid carbon that is sequestered at depth. For the zooplankton that survive overwintering, their upward migration during the spring returns a fraction of the lipid reserves to the surface as nonrespired carbon, with losses attributed to predation by deep-dwelling predators, disease, starvation, and other sources of mortality generally not accounted for. Similar to the
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more nutrient-poor surface waters. Climate change alters the timing of the spring bloom by promoting an earlier or later ice melt. Warmer waters could lead to weaker stratification, meaning the density differences between the first and second layer of the ocean are increasing due to an increased flux of freshwater from ice melt. Typically, the amount of total annual primary productivity in the
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whales. Copepods can account for about 70–90% of total zooplankton biomass, depending on region. Additionally, their eggs are a main source of food for commercially important fish stocks. The copepod eggs are buoyant and will rise to the sea surface, but are susceptible to predation by fish and other organisms. Copepods also provide the
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in arctic and subarctic regions fish for cod, salmon, crab, groundfish, and pollock depend on this energy-rich zooplankton as food. In 2017, the highest value of commercial fish species for the US was salmon ($ 688 million), crabs ($ 610 million), shrimp ($ 531 million), scallops ($ 512 million), and
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found throughout the world's oceans. This is primarily due to zooplankton in their copepodite stages releasing an excessive amount of nitrogen and phosphorus from excretion back into the surface. Thus, the production, transport, and metabolism of lipid carbon during overwintering do not contribute to
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Although the sequestration of marine carbon is a primary outcome of the biological pump, the recycling of nutrients such as N and P in organic matter plays a comparatively important role in maintaining the processes that facilitate this carbon export without removing nutrients for primary production.
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contribute to a significant amount of this POC being attenuated as it sinks below the thermocline (near overwintering depths of ~1000 m). Furthermore, the remaining quantity of carbon in the North Atlantic from the export of POC below the thermocline has been calculated (2–8 g C m y) to be comparable
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The 2015 paper by Jónasdóttir et al., marked the first comprehensive accounting for the amount of carbon sequestration resulting from the movement of lipids by vertically migrating zooplankton during their overwintering diapuse. Although only elucidating the impact of one particular species, in this
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Other climate change factors to consider that might influence these lipid-rich copepods are shifts of current systems, storm activity and sea-ice cover. In some regions of the arctic, specifically the Bering Sea, studies have forecasted a decrease in storms due to warming. This impacts the mixing of
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spp. are not only important for moving carbon out of the photic zone and into the deep ocean, but these lipid-rich organisms play a critical role in the success of many marine species that depend on them as food. They comprise the majority of diets for fishes, seabirds and even large mammals such as
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This data image shows the monthly average sea surface temperature for May 2015. Between 2013 and 2016, a large mass of unusually warm ocean water--nicknamed the blob--dominated the North Pacific, indicated here by red, pink, and yellow colors signifying temperatures as much as three degrees Celsius
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through the export of POC to the deep ocean. Although zooplankton are known to play important roles in the biological pump through grazing and the repackaging of particulate matter, the active transport of seasonally-migrating zooplankton through the lipid pump has not been incorporated into global
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Certain aspects of the lipid pump such as the diapause depth and duration of zooplankton can vary among regions that have different overwintering temperatures and resident community characteristics. There are other subarctic regions that have shown similar carbon export rates to those found in the
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eggs are spawned by females at depth and rise to the surface. Larvae (nauplii) first develop from these eggs, and complete their maturation into an early juvenile (copepodite) within one season, after which they undergo their first overwintering. Copepodite have three stages before maturing to the
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The global estimates of the biological pump have yet to include the elements of the lipid pump which could represent 50–100% of C export that is not accounted for. This is likely due to many observational challenges pertaining to the analysis of these seasonal migrations. As described above, more
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of the ocean in the summertime. Stratification leaves nutrient-rich water on the bottom and nutrient-poor water on the top due to an increase in freshwater from the ice. However, in the wintertime, this region of the world experiences an increase in storms that bring nutrient-rich waters into the
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combine to limit the pelagic primary production. Consequently, the food supply fades toward fall, and overwintering diapause initiates. These copepods migrate to deeper waters with accumulated lipid reserves for overwintering. The overwintering diapause stages remain in deeper waters with limited
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In the Arctic and Antarctic environments, the productive season is typically short and certain copepods species vertically migrate during overwintering diapause. During the productive seasons of spring and summer, younger developmental stages of these copepods usually thrive in food-rich, warmer,
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during seasonal migrations which further explains why the lipid pump has yet to become incorporated into estimates of the global carbon export flux. These observations can be challenging to make given the remote locations they are conducted in and the harsh, deep sampling conditions, but these
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spp. metabolic processes during diapause is required. The importance of diapause timing with spring plankton blooms is well-established, suggesting that there is potential for additional population impacts as a result of climate change, which would further reverberate throughout the ecosystem.
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spp. may deviate in size but their basic physical structure remains constant across different overwintering stages and between different copepod species. The only significant taxonomic difference is the number of segments on the tail across developmental stage CIII and older (CIV, CV). With an
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spp. suggest the possible importance of the lipid pump in global carbon cycling by contributing an estimated 50–100% of carbon sequestration to the biological pump. Subsequent research has underscored this significance as estimates that attempt to more accurately account for the mortality and
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spp. in some ocean basins. In addition to potential ecosystem impacts due to the large number of species that rely on copepods as a major constituent of their diets, there may be implications for oceanic carbon sequestration from consequent changes in the magnitude of the lipid pump due to
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a net consumption or removal of essential nutrients in the surface ocean, which is unlike many components of the biological pump. This process creates what is known as a "lipid shunt" in the biological pump, as the carbon sequestration of the lipid pump is decoupled from nutrient removal.
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are the most dominant zooplankton species found in the Arctic Ocean at similar latitudes, and they contribute to a 3.1 g C m y flux of lipid carbon below 100 m during overwintering. A slightly higher maximum flux in lipid carbon (2–4.3 g C m y) below 150 m was observed in the subarctic
1416:, and will likely have negative effects on whales and other components of the food web that are inextricably tied to copepods. The impact of diapause and variation in seasonal productivity was not explicitly included as increasing model complexity and more accurate accounting for 1136:. While high NAO index values indicate a net flow of Atlantic water to the northeast and into the Norwegian Sea, low NAO index values indicate a reduced Atlantic water inflow into the Nordic Seas. In the Northwestern Atlantic, positive trends in the abundances of 1945:
Sanders, Richard; Henson, Stephanie A.; Koski, Marja; De La Rocha, Christina L.; Painter, Stuart C.; Poulton, Alex J.; Riley, Jennifer; Salihoglu, Baris; Visser, Andre; Yool, Andrew; Bellerby, Richard (2014). "The Biological Carbon Pump in the North Atlantic".
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the water column that brings nutrient-rich water upwards. Copepods consume primary producers that require nutrients to survive. Limiting the amount of nutrients in the water column could decrease the abundance of these primary producers and subsequently reduce
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spp. reside, melting ice caps and timing of the spring phytoplankton bloom could have implications for copepod density, distribution and timing of return from overwintering. A phytoplankton bloom occurs in the spring in arctic and subarctic environments when
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in an oil sac, which can account for up to 60% of an individual's dry weight. Calanus spp. accumulate these lipids while feeding closer to the ocean surface during the spring and summer months, aligning with phytoplankton blooms. Early in the growing season,
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outcome of isomorphism, dry weight (d ) and prosome length (p ) can be scaled as they are related as d = cp, where c is a coefficient. Observations identify the relationship between dry weight and prosome length with a coefficient between 3.3 and 3.5 for
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One key difference between the lipid pump and biological pump is that the ratios of nutrients such as nitrogen and phosphorus relative to carbon are minimal or zero in lipids, whereas the exported POC in the biological pump retains the standard
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spp. (i.e., dry weight, prosome length, lipid content, and carbon content) are always changing, varying between different regions, temporally, and across life stages. Based on isomorphism, or the similarity in form or structure of organisms,
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back to the surface environment, and thus are not removed from the surface mixed layer of the ocean. This means that the carbon transported by the lipid pump does not limit the availability of essential nutrients in the ocean surface.
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Hunt Jr, George L.; Stabeno, Phyllis; Walters, Gary; Sinclair, Elizabeth; Brodeur, Richard D.; Napp, Jeffery M.; Bond, Nicholas A. (2002-12-01). "Climate change and control of the southeastern Bering Sea pelagic ecosystem".
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in the North Atlantic (1–4 g C m y) through the lipid pump. Therefore, the lipid pump may contribute 50–100% of C sequestration to the biological pump as net transport that has not been included in its current estimates.
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Kobari, Toru; Steinberg, Deborah K.; Ueda, Ai; Tsuda, Atsushi; Silver, Mary W.; Kitamura, Minoru (2008). "Impacts of ontogenetically migrating copepods on downward carbon flux in the western subarctic Pacific Ocean".
1214:. The rates or magnitude of these processes may slightly vary due to characteristic differences between these subpolar regions, which have largely been under-studied relative to their contributions to the lipid pump. 726:
to continue. In the Biological Pump, nutrient removal is always coupled to carbon sequestration; primary production is limited as carbon and nutrients are transported to depth together in the form of organic matter.
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Hunt, George L. Jr; Coyle, Kenneth O.; Eisner, Lisa B.; Farley, Edward V.; Heintz, Ron A.; Mueter, Franz; Napp, Jeffrey M.; Overland, James E.; Ressler, Patrick H.; Salo, Sigrid; Stabeno, Phyllis J. (2011-07-01).
1477:, the dynamics of the lipid shunt causes uncertainty in observational methods of the lipid pump when comparing its efficiency to that of the biological pump. Additionally, large zooplankton usually avoid 1387:
Changes in the water masses and temperature could have a direct effect on the zooplankton's vertical migration. The distribution of the zooplankton in the water column is controlled by the currents. The
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excrete waste, that waste falls to the sea floor and organisms on the sea floor compete for the pellets as food. The role of copepods in the food web is crucially intertwined amongst other organisms.
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Jensen, Maj Holst; Nielsen, Torkel Gissel; Dahllöf, Ingela (2008-04-28). "Effects of pyrene on grazing and reproduction of Calanus finmarchicus and Calanus glacialis from Disko Bay, West Greenland".
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and a daytime descent to deeper waters. A relatively unique variation of this form is the twilight DVM, where the ascent happens during dusk and the descent around midnight (i.e., midnight sinking).
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species specifically, the start of reproduction is linked to the start of the spring bloom. Thus, changes in the timing of the spring bloom would directly influence the reproductive capabilities of
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to adapt to the seasonal variability in food availability in ocean basins. Changes in the timing or length of high food periods are likely to negatively impact the distribution and abundance of
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effects by lowering the abundance of larger species and increasing the amount of lipid-rich copepods and even paving way for other species to consume them. Under warming ocean conditions,
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spp. are in diapause could result in a reduction in the abundance of the copepods in the Norwegian Sea. Since the lipid pump is controlled through the movement of copepods, particularly
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spp. biogenergetics are allocated to reproduction, feeding and growth, but eventually shift to the production of lipids to provide energy during diapause. These lipids take the form of
2283:"Investigations on the ecology of Calanus spp. in the Labrador Sea. I. Relationship between the phytoplankton bloom and reproduction and development of Calanus finmarchicus in spring" 730:
The contribution of the lipid pump to the sequestering of carbon in the deeper waters of the ocean can be substantial: the carbon transported below 1,000 metres (3,300 ft) by
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of the eastern shelf of North America forecasts lower abundance of copepods. The decrease in favorable environmental conditions is expected to decrease the size and density of
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Through the seasonal vertical migration of zooplankton, the lipid pump creates a net difference between lipids transported to the deep during the fall (when zooplankton enter
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melts, allowing an increase in light to penetrate deeper into the water column, thus supporting photosynthesis. An input of freshwater from the sea ice melting increases the
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oceans, and previously understood to be the most significant contributor to the active export of carbon as a result of zooplankton migration. The most common form is the
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a day. Each copepod measures about 2–4 millimetres long which is about the size of a grain of rice and they weigh, on average, between 1.0274 and 1.0452 g cm. A loss in
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spp. have suggested similar, although regionally variable, magnitudes of carbon export from the lipid pump. Overwintering diapause is an ecological strategy to enable
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rely on copepods as their primary prey in order to meet their nutritional needs. To meet the right whale's energetic requirements they need about 500 kg of
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spp., impacts of climate change that affect copepod abundance or seasonal migration will directly impact the lipid pump and carbon export to the deep ocean.
1343:(five degrees Fahrenheit) higher than average. Data are from the NASA Multi-scale Ultra-high Resolution Sea Surface Temperature (MUR SST) Analysis product." 925:
physical and physiological activity and ascend back to the near-surface waters and complete the life cycle at the onset of the following productive season.
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A study that utilized climate modeling to simulate the effects of predicted increases in water temperature and salinity as a result of climate change on
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has the potential to affect the right whale's migration, reproduction, and/or ability to successfully nurse their young (only for lactating females).
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Kobari, Toru; Shinada, Akiyoshi; Tsuda, Atsushi (2003). "Functional roles of interzonal migrating mesozooplankton in the western subarctic Pacific".
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near-surface waters, and they rapidly develop and grow. During late summer and fall, grazing pressure, nutrient limitation, and annual variations of
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to prevent premature return to the surface waters. Stored lipids are metabolized at these depths, accounting for approximately 25% of the
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spp. in ocean basins with shorter growth seasons will be increasingly sensitive to the timing of the spring bloom, such as polar regions.
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adaptations in the data collection are needed to better integrate global estimates of the carbon export flux provided by the lipid pump.
1074:. A 6–8 month-long overwintering period can drain a substantial fraction (44–93%) of the stored lipids despite the decreased metabolism. 48: 1330:
associated with a spring bloom is approximately 10–65%, however warmer waters could impact the amount of primary production occurring.
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is estimated to impact the marine environment in a variety of ways. In the arctic and subarctic environments where a vast majority of
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Changes in ocean temperature and salinity due to anthropogenic climate change are also predicted to decrease concentrations of
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out of the surface waters was found to be 29 ± 10 g C m y. However, studies have shown that processes such as consumption and
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and subpolar regions containing seasonally-migrating zooplankton. The role of zooplankton, and particularly copepods, in the
2209:"Projecting the effects of climate change on Calanus finmarchicus distribution within the U.S. Northeast Continental Shelf" 1283: 1360: 1356: 2103:"Zooplankton vertical migration and the active transport of dissolved organic and inorganic nitrogen in the Sargasso Sea" 1367:
is to be expected. Egg production and hatching success may also be affected with increasing sea surface temperatures and
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Visser, Andre W.; Grønning, Josephine; Jónasdóttir, Sigrún Huld (2017). "Calanus hyperboreus and the lipid pump".
799:) and what returns to the surface during the spring, resulting in the sequestration of lipid carbon at depth. The 573: 193: 634: 2482:"Potential contribution that the copepod Neocalanus tonsus makes to downward carbon flux in the Southern Ocean" 695: 548: 476: 471: 454: 299: 160: 58: 2481: 2662:
Schumacher, J.D.; Bond, N.; Brodeur, Richard; Livingston, Patricia; Napp, J.M.; Stabeno, P.J. (2003-01-01),
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abundance was substantially diminished when temporal autocorrelation and detrending analyses were involved.
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Kristiansen, Inga; Gaard, Eilif; Hátún, Hjálmar; Jónasdóttir, Sigrún; Ferreira, A. Sofia A. (2016-05-01).
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and positive NAO forcing with a lag of one or two years. However, the influence of the NAO in explaining
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Siegel, D. A.; Buesseler, K. O.; Doney, S. C.; Sailley, S. F.; Behrenfeld, M. J.; Boyd, P. W. (2014).
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Jónasdóttir, Sigrún Huld; Visser, André W.; Richardson, Katherine; Heath, Michael R. (2015-09-29).
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via the lipid pump is therefore decoupled from nutrient removal, allowing carbon uptake by oceanic
439: 255: 110: 2916:"Global assessment of ocean carbon export by combining satellite observations and food-web models" 1000:(hibernation) strategy, and its life-history will be described in more detail as a representative 2953: 2896: 2841: 2783: 2724: 2390: 2083: 2018: 1710: 1051: 901: 777: 723: 518: 286: 270: 265: 188: 1338: 967:. Studies attempting to quantify the lipid pump have primarily focused on the cousin species of 2145:"Zooplankton respiration and the export of carbon at depth in the Amundsen Gulf (Arctic Ocean)" 2888: 2880: 2833: 2775: 2716: 2608: 2563: 2382: 2312: 2254: 2236: 2174: 2075: 2067: 2010: 1924: 1916: 1872: 1822: 1776: 1768: 1702: 1614: 973: 936: 749: 449: 228: 63: 2943: 2935: 2872: 2825: 2767: 2706: 2598: 2553: 2493: 2459: 2420: 2372: 2362: 2302: 2244: 2228: 2164: 2122: 2059: 2002: 1963: 1908: 1862: 1812: 1760: 1694: 1604: 1594: 839: 415: 309: 304: 1171:
temperate North Atlantic (1–4 g C m y) via seasonally-migrating zooplankton. For instance,
892:(hibernation) occurs on an annual time-scale and enables zooplankton species, particularly 32: 2039:"High-latitude controls of thermocline nutrients and low latitude biological productivity" 1495: 1062:
migrate downward, with to depths varying from 600 to 3000m, but with the requirement that
1022: 800: 706: 543: 410: 343: 331: 260: 134: 1764: 2931: 2821: 2763: 2702: 2455: 2358: 2298: 2224: 2160: 2118: 2102: 2055: 1998: 1959: 1756: 1690: 2249: 2208: 1609: 1582: 1364: 1309: 1270: 1203: 964: 909: 858: 835: 769: 699: 691: 553: 523: 503: 444: 358: 241: 236: 97: 2771: 2516: 2424: 2282: 2126: 1583:"Seasonal copepod lipid pump promotes carbon sequestration in the deep North Atlantic" 1392:
spp. use the water column for their vertical migration. Changes to the currents while
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deficient compounds essential for cellular structures. This lipid carbon enters the
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community with food via sinking fecal pellets, meaning that as fish and smaller
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Steinberg, Deborah K.; Goldthwait, Sarah A.; Hansell, Dennis A. (2002-08-01).
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Climate change in the southeastern Bering Sea and some consequences for biota
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are abundantly distributed copepods, particularly in the polar and temperate
2915: 2687:"Trend and Variability in Global Upper-Ocean Stratification Since the 1960s" 2036:
Sarmiento, J. L.; Gruber, N.; Brzezinski, M. A.; Dunne, J. P. (2004-01-01).
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has a direct impact on the endangered right whales of the North Atlantic.
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are the primary contributors to the lipid pump, whereas, the subantarctic
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Parent, Genevieve J.; Plourde, Stephane; Turgeon, Julie (2011-11-01).
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individual can go through multiple overwintering periods. Positively
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contributing to a lipid carbon flux of 1.7–9.3 g C m y out of the
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like nitrogen and phosphorus that are consumed in excess must be
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almost equals that transported below the same depth annually by
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Grieve, Brian D.; Hare, Jon A.; Saba, Vincent S. (2017-07-24).
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is a well-studied phenomenon, widespread in the temperate and
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Head, E. J. H.; Harris, L. R.; Campbell, R. W. (2000-02-28).
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The biological pump transports 1–4 g C m y of POC below the
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is copepods. With warming oceans and increasing melting of
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Deep Sea Research Part II: Topical Studies in Oceanography
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Knutsen, Tor; Melle, Webjorn; Calise, Lucio (2001-08-01).
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Deep Sea Research Part II: Topical Studies in Oceanography
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spp. are generally expected to experience mortality after
2861:"Marine viruses — major players in the global ecosystem" 1739:
Steinberg, Deborah K.; Landry, Michael R. (2017-01-03).
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Global distribution of particulate organic carbon (POC).
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Deep Sea Research Part I: Oceanographic Research Papers
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With a life cycle of two to six years on average, each
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(2019). 1274:Relationship of organisms in an arctic system. 1033:Lipids are stored by all copepodite and adult 662:from the ocean's surface to deeper waters via 635: 8: 1287:Pollock fishing vessels in an Alaskan port. 1120:, a primary long-term forcing that affects 642: 628: 15: 2947: 2710: 2602: 2557: 2376: 2366: 2306: 2248: 2168: 1866: 1816: 1608: 1598: 1439:respiration rates of other overwintering 2685:Yamaguchi, Ryohei; Suga, Toshio (2019). 1741:"Zooplankton and the Ocean Carbon Cycle" 1334:Reproduction and changes to the food web 1282: 1269: 1226: 1159: 819: 2691:Journal of Geophysical Research: Oceans 2579: 2577: 2336: 2334: 2332: 2330: 2328: 2326: 2149:Journal of Geophysical Research: Oceans 2143:Darnis, Gérald; Fortier, Louis (2012). 1512: 579:Territorialisation of carbon governance 23: 2799: 2797: 2744: 2742: 2740: 2738: 2657: 2655: 2653: 2651: 2649: 2647: 2626: 2624: 2622: 2511: 2509: 2507: 2276: 2274: 2272: 2270: 2268: 2202: 2200: 2198: 2196: 2194: 2192: 2190: 2188: 1672: 1670: 1668: 1666: 1664: 1662: 1660: 1658: 1656: 1654: 1652: 1650: 1648: 881:DVM, a night-time ascent to the upper 2480:Bradford-Grieve, J. M. (2001-09-01). 2475: 2473: 2436: 2434: 2406: 2404: 2138: 2136: 1894: 1892: 1890: 1888: 1886: 1844: 1842: 1840: 1838: 1836: 1794: 1792: 1790: 1646: 1644: 1642: 1640: 1638: 1636: 1634: 1632: 1630: 1628: 1576: 1574: 1572: 1570: 1568: 1566: 1564: 1562: 1560: 1558: 1556: 1554: 1552: 1550: 1548: 1546: 1544: 1542: 1540: 1538: 1536: 756:is crucial to the survival of higher 584:Total Carbon Column Observing Network 7: 1940: 1938: 1765:10.1146/annurev-marine-010814-015924 1734: 1732: 1730: 1728: 1726: 1724: 1534: 1532: 1530: 1528: 1526: 1524: 1522: 1520: 1518: 1516: 1251:Copepod abundance, specifically the 1184:and was primarily attributed to the 1190:genus of copepods. In these areas, 702:from the mortality of zooplankton. 1355:and alter the food chain from the 812:estimates of the biological pump. 760:organisms whose primary source of 14: 1231:Planktonic relationships to fish. 1029:Lipid accumulation and metabolism 825:Components of the biological pump 807:taken up in the surface ocean by 2754:. Ecology of the SE Bering Sea. 1291:Many commercial and subsistence 1082:The physical characteristics of 609: 608: 31: 2640:. Washington: Govt. Print. Off. 1745:Annual Review of Marine Science 2634:Fisheries of the United States 2591:ICES Journal of Marine Science 2287:Marine Ecology Progress Series 1855:ICES Journal of Marine Science 1104:Relationships between NAO and 791:Lipid pump vs. biological pump 544:Climate reconstruction proxies 1: 2772:10.1016/S0967-0645(02)00321-1 2425:10.1016/S0079-6611(03)00102-2 2127:10.1016/S0967-0637(02)00037-7 1913:10.1016/j.aquatox.2008.01.005 1050:, energy-rich compounds like 947:ranging from 4–7 millimeters. 871:Diel Vertical Migration (DVM) 843:to the seasonal migration of 816:Comparison between net fluxes 705:Compared to the more general 2920:Global Biogeochemical Cycles 2546:Journal of Plankton Research 2486:Journal of Plankton Research 1968:10.1016/j.pocean.2014.05.005 1805:Journal of Plankton Research 1140:spp. correspond with higher 1124:spp. and its habitat is the 514:Carbonate compensation depth 179:Particulate inorganic carbon 2865:Nature Reviews Microbiology 2521:repository.library.noaa.gov 1981:Falkowski, Paul G. (1997). 1464:overwintering zooplankton. 1257:North Atlantic right whales 1017:adult stages. While female 2999: 2859:Suttle, Curtis A. (2007). 2464:10.1016/j.dsr2.2008.04.016 2347:Limnology and Oceanography 2233:10.1038/s41598-017-06524-1 1679:Limnology and Oceanography 1126:North Atlantic Oscillation 1116:In the North Atlantic and 746:particulate organic carbon 569:Carbon capture and storage 173:Particulate organic carbon 167:Dissolved inorganic carbon 910:match-mismatch hypothesis 698:of lipid reserves and as 574:Carbon cycle re-balancing 2413:Progress in Oceanography 1948:Progress in Oceanography 1384:spp. abundance as well. 1142:sea surface temperatures 1078:Physical characteristics 674:. Lipids are a class of 549:Carbon-to-nitrogen ratio 509:Carbonate–silicate cycle 477:Carbon dioxide clathrate 472:Clathrate gun hypothesis 300:Net ecosystem production 161:Dissolved organic carbon 2559:10.1093/plankt/23.8.859 2498:10.1093/plankt/23.9.963 1600:10.1073/pnas.1512110112 559:Deep Carbon Observatory 19:Part of a series on the 2983:Carbon dioxide removal 2604:10.1093/icesjms/fsr036 1868:10.1093/icesjms/fsv222 1344: 1304:Climate change impacts 1288: 1275: 1232: 1206:consists primarily of 1167: 948: 827: 785:global oceanic systems 379:Continental shelf pump 155:Total inorganic carbon 121:Satellite measurements 2978:Chemical oceanography 1818:10.1093/plankt/fbr072 1341: 1286: 1273: 1230: 1163: 1100:spp. carbon content. 939: 823: 670:vertically migratory 564:Global Carbon Project 295:Ecosystem respiration 2940:10.1002/2013GB004743 2712:10.1029/2019JC015439 2450:(14–15): 1648–1660. 2170:10.1029/2011JC007374 1501:Oceanic carbon cycle 1481:instruments such as 1223:Role in the food web 1156:Regional differences 1072:basal metabolic rate 902:primary productivity 720:Carbon sequestration 393:Carbon sequestration 149:Total organic carbon 2932:2014GBioC..28..181S 2877:10.1038/nrmicro1750 2822:1979Natur.282..677E 2764:2002DSRII..49.5821H 2703:2019JGRC..124.8933Y 2456:2008DSRII..55.1648K 2359:2019LimOc..64.2044W 2299:2000MEPS..193...53H 2225:2017NatSR...7.6264G 2161:2012JGRC..117.4013D 2119:2002DSRI...49.1445S 2064:10.1038/nature02127 2056:2004Natur.427...56S 1999:1997Natur.387..272F 1960:2014PrOce.129..200S 1757:2017ARMS....9..413S 1691:2017LimOc..62.1155V 1593:(39): 12122–12126. 1369:ocean acidification 1052:omega-3 fatty acids 943:Calanus hyperboreus 440:Atmospheric methane 406:Soil carbon storage 256:Reverse Krebs cycle 111:Ocean acidification 2308:10.3354/meps193053 2213:Scientific Reports 1901:Aquatic Toxicology 1345: 1289: 1276: 1235:The zooplanktonic 1233: 1218:Ecological impacts 1168: 949: 898:seasonal variation 896:spp., to adapt to 828: 724:primary production 519:Great Calcite Belt 467:Aerobic production 287:Carbon respiration 229:Metabolic pathways 189:Primary production 2816:(5740): 677–680. 2758:(26): 5821–5853. 2697:(12): 8933–8948. 2368:10.1002/lno.11168 1993:(6630): 272–275. 1811:(11): 1654–1665. 1699:10.1002/lno.10492 1468:Future directions 1066:settle below the 974:Calanus glacialis 652: 651: 450:Methane emissions 106:In the atmosphere 2990: 2962: 2961: 2951: 2911: 2905: 2904: 2856: 2850: 2849: 2830:10.1038/282677a0 2801: 2792: 2791: 2746: 2733: 2732: 2714: 2682: 2676: 2675: 2674: 2673: 2668:, pp. 17–40 2659: 2642: 2641: 2639: 2628: 2617: 2616: 2606: 2597:(6): 1230–1243. 2581: 2572: 2571: 2561: 2537: 2531: 2530: 2528: 2527: 2513: 2502: 2501: 2477: 2468: 2467: 2438: 2429: 2428: 2419:(3–4): 279–298. 2408: 2399: 2398: 2380: 2370: 2353:(5): 2044–2058. 2338: 2321: 2320: 2310: 2278: 2263: 2262: 2252: 2204: 2183: 2182: 2172: 2140: 2131: 2130: 2113:(8): 1445–1461. 2098: 2092: 2091: 2041: 2033: 2027: 2026: 2007:10.1038/387272a0 1978: 1972: 1971: 1942: 1933: 1932: 1896: 1881: 1880: 1870: 1861:(5): 1319–1329. 1846: 1831: 1830: 1820: 1796: 1785: 1784: 1736: 1719: 1718: 1685:(3): 1155–1165. 1674: 1623: 1622: 1612: 1602: 1578: 1425:Key implications 1404:Climate modeling 1279:Economic impacts 1253:C. finmarchicus, 840:remineralization 666:associated with 644: 637: 630: 617: 612: 611: 416:pelagic sediment 310:Soil respiration 305:Photorespiration 35: 16: 2998: 2997: 2993: 2992: 2991: 2989: 2988: 2987: 2968: 2967: 2966: 2965: 2913: 2912: 2908: 2871:(10): 801–812. 2858: 2857: 2853: 2803: 2802: 2795: 2748: 2747: 2736: 2684: 2683: 2679: 2671: 2669: 2661: 2660: 2645: 2637: 2630: 2629: 2620: 2583: 2582: 2575: 2539: 2538: 2534: 2525: 2523: 2515: 2514: 2505: 2479: 2478: 2471: 2440: 2439: 2432: 2410: 2409: 2402: 2340: 2339: 2324: 2280: 2279: 2266: 2206: 2205: 2186: 2142: 2141: 2134: 2100: 2099: 2095: 2050:(6969): 56–60. 2035: 2034: 2030: 1980: 1979: 1975: 1944: 1943: 1936: 1898: 1897: 1884: 1848: 1847: 1834: 1798: 1797: 1788: 1738: 1737: 1722: 1676: 1675: 1626: 1580: 1579: 1514: 1509: 1496:Biological pump 1492: 1470: 1432:C. finmarchicus 1427: 1410:C. finmarchicus 1406: 1377: 1353:C. finmarchicus 1349:C. finmarchicus 1336: 1306: 1281: 1265:C. finmarchicus 1261:C. finmarchicus 1225: 1220: 1158: 1010:C. hyperboreous 994:C. hyperboreous 970:C. finmarchicus 934: 868: 859:Redfield ratios 854: 845:C. finmarchicus 818: 806: 801:biological pump 793: 707:biological pump 648: 607: 600: 599: 598: 538: 530: 529: 528: 493: 483: 482: 481: 434: 424: 423: 422: 411:Marine sediment 395: 385: 384: 383: 344:Solubility pump 332:Biological pump 326: 316: 315: 314: 289: 279: 278: 277: 261:Carbon fixation 246: 231: 221: 220: 219: 200: 184: 137: 135:Forms of carbon 127: 126: 125: 100: 90: 89: 88: 43: 12: 11: 5: 2996: 2994: 2986: 2985: 2980: 2970: 2969: 2964: 2963: 2926:(3): 181–196. 2906: 2851: 2793: 2734: 2677: 2643: 2618: 2573: 2552:(8): 859–873. 2532: 2503: 2492:(9): 963–975. 2469: 2430: 2400: 2322: 2264: 2184: 2132: 2093: 2028: 1973: 1934: 1882: 1832: 1786: 1751:(1): 413–444. 1720: 1624: 1511: 1510: 1508: 1505: 1504: 1503: 1498: 1491: 1488: 1483:sediment traps 1469: 1466: 1426: 1423: 1414:C. finmarchius 1405: 1402: 1376: 1375:Physical ocean 1373: 1365:prey switching 1335: 1332: 1323:stratification 1310:climate change 1308:Anthropogenic 1305: 1302: 1280: 1277: 1224: 1221: 1219: 1216: 1204:Southern Ocean 1177:C. hyperboreus 1157: 1154: 1094:C. hyperboreus 989:C. hyperboreus 965:North Atlantic 933: 927: 867: 864: 853: 850: 836:North Atlantic 817: 814: 804: 792: 789: 781:marine mammals 770:climate change 700:organic matter 692:carbon dioxide 650: 649: 647: 646: 639: 632: 624: 621: 620: 619: 618: 602: 601: 597: 596: 591: 586: 581: 576: 571: 566: 561: 556: 554:Deep biosphere 551: 546: 540: 539: 536: 535: 532: 531: 527: 526: 524:Redfield ratio 521: 516: 511: 506: 504:Nutrient cycle 501: 495: 494: 491:Biogeochemical 489: 488: 485: 484: 480: 479: 474: 469: 464: 463: 462: 457: 447: 445:Methanogenesis 442: 436: 435: 430: 429: 426: 425: 421: 420: 419: 418: 408: 403: 397: 396: 391: 390: 387: 386: 382: 381: 376: 371: 366: 361: 359:Microbial loop 356: 351: 346: 341: 340: 339: 328: 327: 322: 321: 318: 317: 313: 312: 307: 302: 297: 291: 290: 285: 284: 281: 280: 276: 275: 274: 273: 268: 258: 253: 247: 245: 244: 242:Chemosynthesis 239: 237:Photosynthesis 233: 232: 227: 226: 223: 222: 218: 217: 212: 207: 201: 199: 198: 197: 196: 185: 183: 182: 176: 170: 164: 158: 152: 146: 139: 138: 133: 132: 129: 128: 124: 123: 118: 113: 108: 102: 101: 98:Carbon dioxide 96: 95: 92: 91: 87: 86: 81: 76: 71: 66: 61: 56: 51: 45: 44: 41: 40: 37: 36: 28: 27: 21: 20: 13: 10: 9: 6: 4: 3: 2: 2995: 2984: 2981: 2979: 2976: 2975: 2973: 2959: 2955: 2950: 2945: 2941: 2937: 2933: 2929: 2925: 2921: 2917: 2910: 2907: 2902: 2898: 2894: 2890: 2886: 2882: 2878: 2874: 2870: 2866: 2862: 2855: 2852: 2847: 2843: 2839: 2835: 2831: 2827: 2823: 2819: 2815: 2811: 2807: 2800: 2798: 2794: 2789: 2785: 2781: 2777: 2773: 2769: 2765: 2761: 2757: 2753: 2745: 2743: 2741: 2739: 2735: 2730: 2726: 2722: 2718: 2713: 2708: 2704: 2700: 2696: 2692: 2688: 2681: 2678: 2667: 2666: 2658: 2656: 2654: 2652: 2650: 2648: 2644: 2636: 2635: 2631:NOAA (2017). 2627: 2625: 2623: 2619: 2614: 2610: 2605: 2600: 2596: 2592: 2588: 2580: 2578: 2574: 2569: 2565: 2560: 2555: 2551: 2547: 2543: 2536: 2533: 2522: 2518: 2512: 2510: 2508: 2504: 2499: 2495: 2491: 2487: 2483: 2476: 2474: 2470: 2465: 2461: 2457: 2453: 2449: 2445: 2437: 2435: 2431: 2426: 2422: 2418: 2414: 2407: 2405: 2401: 2396: 2392: 2388: 2384: 2379: 2374: 2369: 2364: 2360: 2356: 2352: 2348: 2344: 2337: 2335: 2333: 2331: 2329: 2327: 2323: 2318: 2314: 2309: 2304: 2300: 2296: 2292: 2288: 2284: 2277: 2275: 2273: 2271: 2269: 2265: 2260: 2256: 2251: 2246: 2242: 2238: 2234: 2230: 2226: 2222: 2218: 2214: 2210: 2203: 2201: 2199: 2197: 2195: 2193: 2191: 2189: 2185: 2180: 2176: 2171: 2166: 2162: 2158: 2154: 2150: 2146: 2139: 2137: 2133: 2128: 2124: 2120: 2116: 2112: 2108: 2104: 2097: 2094: 2089: 2085: 2081: 2077: 2073: 2069: 2065: 2061: 2057: 2053: 2049: 2045: 2040: 2032: 2029: 2024: 2020: 2016: 2012: 2008: 2004: 2000: 1996: 1992: 1988: 1984: 1977: 1974: 1969: 1965: 1961: 1957: 1953: 1949: 1941: 1939: 1935: 1930: 1926: 1922: 1918: 1914: 1910: 1907:(2): 99–107. 1906: 1902: 1895: 1893: 1891: 1889: 1887: 1883: 1878: 1874: 1869: 1864: 1860: 1856: 1852: 1845: 1843: 1841: 1839: 1837: 1833: 1828: 1824: 1819: 1814: 1810: 1806: 1802: 1795: 1793: 1791: 1787: 1782: 1778: 1774: 1770: 1766: 1762: 1758: 1754: 1750: 1746: 1742: 1735: 1733: 1731: 1729: 1727: 1725: 1721: 1716: 1712: 1708: 1704: 1700: 1696: 1692: 1688: 1684: 1680: 1673: 1671: 1669: 1667: 1665: 1663: 1661: 1659: 1657: 1655: 1653: 1651: 1649: 1647: 1645: 1643: 1641: 1639: 1637: 1635: 1633: 1631: 1629: 1625: 1620: 1616: 1611: 1606: 1601: 1596: 1592: 1588: 1584: 1577: 1575: 1573: 1571: 1569: 1567: 1565: 1563: 1561: 1559: 1557: 1555: 1553: 1551: 1549: 1547: 1545: 1543: 1541: 1539: 1537: 1535: 1533: 1531: 1529: 1527: 1525: 1523: 1521: 1519: 1517: 1513: 1506: 1502: 1499: 1497: 1494: 1493: 1489: 1487: 1484: 1480: 1476: 1467: 1465: 1462: 1458: 1454: 1450: 1446: 1442: 1437: 1433: 1424: 1422: 1419: 1415: 1411: 1403: 1401: 1399: 1395: 1391: 1385: 1383: 1374: 1372: 1370: 1366: 1362: 1358: 1354: 1350: 1340: 1333: 1331: 1329: 1324: 1320: 1315: 1311: 1303: 1301: 1299: 1294: 1285: 1278: 1272: 1268: 1266: 1262: 1258: 1254: 1249: 1247: 1246:invertebrates 1243: 1238: 1229: 1222: 1217: 1215: 1213: 1212:euphotic zone 1209: 1205: 1201: 1197: 1193: 1192:N. flemingeri 1189: 1188: 1183: 1182:North Pacific 1178: 1174: 1166: 1162: 1155: 1153: 1151: 1147: 1143: 1139: 1135: 1134:Icelandic Low 1131: 1127: 1123: 1119: 1114: 1113: 1111: 1107: 1101: 1099: 1095: 1090: 1085: 1080: 1079: 1075: 1073: 1069: 1065: 1061: 1057: 1053: 1049: 1045: 1040: 1036: 1031: 1030: 1026: 1024: 1020: 1015: 1011: 1007: 1003: 999: 995: 991: 990: 985: 984: 983:helgolandicus 981: 976: 975: 971: 966: 962: 958: 954: 953: 946: 944: 938: 931: 928: 926: 923: 917: 915: 911: 907: 903: 899: 895: 891: 886: 884: 880: 876: 872: 865: 863: 860: 851: 849: 846: 841: 837: 833: 826: 822: 815: 813: 810: 809:phytoplankton 802: 798: 790: 788: 786: 782: 779: 775: 771: 767: 763: 759: 758:trophic level 755: 751: 747: 743: 739: 738: 734:of the genus 733: 728: 725: 721: 716: 712: 708: 703: 701: 697: 693: 689: 685: 681: 677: 673: 669: 668:overwintering 665: 661: 657: 645: 640: 638: 633: 631: 626: 625: 623: 622: 616: 606: 605: 604: 603: 595: 592: 590: 587: 585: 582: 580: 577: 575: 572: 570: 567: 565: 562: 560: 557: 555: 552: 550: 547: 545: 542: 541: 534: 533: 525: 522: 520: 517: 515: 512: 510: 507: 505: 502: 500: 499:Marine cycles 497: 496: 492: 487: 486: 478: 475: 473: 470: 468: 465: 461: 458: 456: 453: 452: 451: 448: 446: 443: 441: 438: 437: 433: 428: 427: 417: 414: 413: 412: 409: 407: 404: 402: 399: 398: 394: 389: 388: 380: 377: 375: 372: 370: 367: 365: 362: 360: 357: 355: 352: 350: 347: 345: 342: 338: 335: 334: 333: 330: 329: 325: 320: 319: 311: 308: 306: 303: 301: 298: 296: 293: 292: 288: 283: 282: 272: 269: 267: 264: 263: 262: 259: 257: 254: 252: 249: 248: 243: 240: 238: 235: 234: 230: 225: 224: 216: 213: 211: 208: 206: 203: 202: 195: 192: 191: 190: 187: 186: 180: 177: 174: 171: 168: 165: 162: 159: 156: 153: 150: 147: 144: 141: 140: 136: 131: 130: 122: 119: 117: 114: 112: 109: 107: 104: 103: 99: 94: 93: 85: 82: 80: 79:Boreal forest 77: 75: 72: 70: 67: 65: 62: 60: 57: 55: 52: 50: 47: 46: 39: 38: 34: 30: 29: 26: 22: 18: 17: 2923: 2919: 2909: 2868: 2864: 2854: 2813: 2809: 2755: 2751: 2694: 2690: 2680: 2670:, retrieved 2664: 2633: 2594: 2590: 2549: 2545: 2535: 2524:. Retrieved 2520: 2489: 2485: 2447: 2443: 2416: 2412: 2350: 2346: 2290: 2286: 2216: 2212: 2152: 2148: 2110: 2106: 2096: 2047: 2043: 2031: 1990: 1986: 1976: 1951: 1947: 1904: 1900: 1858: 1854: 1808: 1804: 1748: 1744: 1682: 1678: 1590: 1586: 1471: 1460: 1456: 1452: 1448: 1444: 1440: 1435: 1431: 1428: 1417: 1413: 1409: 1407: 1397: 1393: 1389: 1386: 1381: 1378: 1352: 1348: 1346: 1313: 1307: 1290: 1264: 1260: 1252: 1250: 1236: 1234: 1207: 1200:N. plumchrus 1199: 1196:N. cristatus 1195: 1191: 1185: 1176: 1173:C. glacialis 1172: 1169: 1164: 1149: 1145: 1137: 1121: 1115: 1109: 1105: 1103: 1102: 1097: 1093: 1088: 1083: 1081: 1077: 1076: 1064:Calanus spp. 1063: 1059: 1055: 1043: 1038: 1034: 1032: 1028: 1027: 1018: 1009: 1005: 1001: 993: 987: 982: 979: 968: 960: 956: 955: 951: 950: 941: 940:The copepod 929: 918: 913: 906:ocean basins 904:in specific 893: 887: 869: 855: 844: 829: 824: 794: 742:Arctic Ocean 735: 729: 704: 694:produced by 655: 653: 348: 337:Martin curve 324:Carbon pumps 251:Calvin cycle 205:Black carbon 143:Total carbon 84:Geochemistry 25:Carbon cycle 2378:10037/15827 2219:(1): 6264. 1954:: 200–218. 1475:lysis shunt 1130:Azores High 1118:Nordic Seas 1112:populations 1068:thermocline 852:Lipid shunt 832:thermocline 696:respiration 676:hydrocarbon 672:zooplankton 658:sequesters 401:Carbon sink 364:Viral shunt 354:Marine snow 210:Blue carbon 64:Deep carbon 59:Atmospheric 49:Terrestrial 2972:Categories 2672:2021-11-14 2526:2021-11-14 1507:References 1328:Bering Sea 1187:Neocalanus 1048:wax esters 922:irradiance 778:endangered 688:deep ocean 684:phosphorus 656:lipid pump 374:Whale pump 369:Jelly pump 349:Lipid pump 74:Permafrost 42:By regions 2949:1912/6668 2885:1740-1526 2838:1476-4687 2780:0967-0645 2729:213693676 2721:2169-9291 2613:1054-3139 2568:0142-7873 2395:146002715 2387:1939-5590 2317:0171-8630 2293:: 53–73. 2241:2045-2322 2179:2156-2202 2072:0028-0836 2015:1476-4687 1921:0166-445X 1877:1054-3139 1827:0142-7873 1773:1941-1405 1707:1939-5590 1357:bottom-up 1293:fisheries 1208:N. tonsus 879:nocturnal 762:nutrition 750:temperate 711:nutrients 2958:43799803 2893:17853907 2846:42385900 2788:55222333 2259:28740241 2088:52798128 2080:14702082 1929:18291539 1781:27814033 1715:51989153 1619:26338976 1490:See also 1361:top-down 1347:For the 1132:and the 1023:spawning 998:diapause 890:diapause 875:tropical 797:diapause 766:ice caps 754:food web 732:copepods 715:excreted 680:nitrogen 615:Category 2928:Bibcode 2901:4658457 2818:Bibcode 2760:Bibcode 2699:Bibcode 2452:Bibcode 2355:Bibcode 2295:Bibcode 2250:5524788 2221:Bibcode 2157:Bibcode 2115:Bibcode 2052:Bibcode 2023:4326172 1995:Bibcode 1956:Bibcode 1753:Bibcode 1687:Bibcode 1610:4593097 1479:mooring 1461:Calanus 1453:Calanus 1445:Calanus 1441:Calanus 1436:Calanus 1418:Calanus 1398:Calanus 1394:Calanus 1390:Calanus 1382:Calanus 1319:sea ice 1314:Calanus 1298:pollock 1242:benthic 1237:Calanus 1146:Calanus 1138:Calanus 1122:Calanus 1106:Calanus 1098:Calanus 1089:Calanus 1084:Calanus 1056:Calanus 1044:Calanus 1035:Calanus 1019:Calanus 1014:buoyant 1002:Calanus 980:Calanus 957:Calanus 952:Ecology 930:Calanus 914:Calanus 894:Calanus 883:pelagic 768:due to 740:in the 737:Calanus 460:Wetland 432:Methane 215:Kerogen 116:Removal 2956:  2899:  2891:  2883:  2844:  2836:  2810:Nature 2786:  2778:  2727:  2719:  2611:  2566:  2393:  2385:  2315:  2257:  2247:  2239:  2177:  2155:(C4). 2086:  2078:  2070:  2044:Nature 2021:  2013:  1987:Nature 1927:  1919:  1875:  1825:  1779:  1771:  1713:  1705:  1617:  1607:  1430:case, 1198:, and 678:rich, 664:lipids 660:carbon 613:  594:CO2SYS 455:Arctic 194:marine 54:Marine 2954:S2CID 2897:S2CID 2842:S2CID 2784:S2CID 2725:S2CID 2638:(PDF) 2391:S2CID 2084:S2CID 2019:S2CID 1711:S2CID 589:C4MIP 537:Other 181:(PIC) 175:(POC) 169:(DIC) 163:(DOC) 157:(TIC) 151:(TOC) 2889:PMID 2881:ISSN 2834:ISSN 2776:ISSN 2717:ISSN 2609:ISSN 2564:ISSN 2383:ISSN 2313:ISSN 2255:PMID 2237:ISSN 2175:ISSN 2076:PMID 2068:ISSN 2011:ISSN 1925:PMID 1917:ISSN 1873:ISSN 1823:ISSN 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Index

Carbon cycle

Terrestrial
Marine
Atmospheric
Deep carbon
Soil
Permafrost
Boreal forest
Geochemistry
Carbon dioxide
In the atmosphere
Ocean acidification
Removal
Satellite measurements
Forms of carbon
Total carbon
Total organic carbon
Total inorganic carbon
Dissolved organic carbon
Dissolved inorganic carbon
Particulate organic carbon
Particulate inorganic carbon
Primary production
marine
Black carbon
Blue carbon
Kerogen
Metabolic pathways
Photosynthesis

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