Biological Pump
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The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and
seafloor sediments Marine sediment, or ocean sediment, or seafloor sediment, are deposits of insoluble particles that have accumulated on the seafloor. These particles have their origins in soil and rocks and have been transported from the land to the sea, mainly ...
.Sigman DM & GH Haug. 2006. The biological pump in the past. In: Treatise on Geochemistry; vol. 6, (ed.). Pergamon Press, pp. 491-528 In other words, it is a biologically mediated processes which result in the sequestering of carbon in the deep ocean away from the atmosphere and the land. The biological pump is the biological component of the "marine carbon pump" which contains both a physical and biological component. It is the part of the broader
oceanic carbon cycle The oceanic carbon cycle (or marine carbon cycle) is composed of processes that exchange carbon between various pools within the ocean as well as between the atmosphere, Earth interior, and the seafloor. The carbon cycle is a result of many inter ...
responsible for the cycling of
organic matter Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
formed mainly by
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...
during
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
(soft-tissue pump), as well as the cycling of calcium carbonate (CaCO3) formed into shells by certain organisms such as
plankton Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
and
mollusks Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
(carbonate pump). Budget calculations of the biological carbon pump are based on the ratio between
sedimentation Sedimentation is the deposition of sediments. It takes place when particles in suspension settle out of the fluid in which they are entrained and come to rest against a barrier. This is due to their motion through the fluid in response to the ...
(carbon export to the ocean floor) and
remineralization In biogeochemistry, remineralisation (or remineralization) refers to the breakdown or transformation of organic matter (those molecules derived from a biological source) into its simplest Inorganic compound, inorganic forms. These transformations ...
(release of carbon to the atmosphere). The biological pump is not so much the result of a single process, but rather the sum of a number of processes each of which can influence biological pumping. Overall, the pump transfers about 11
gigatonne The tonne ( or ; symbol: t) is a unit of mass equal to 1000 kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton (United States c ...
s of carbon every year into the ocean's interior. This takes carbon out of contact with the atmosphere for several thousand years or longer. An ocean without a biological pump would result in atmospheric carbon dioxide levels about 400 ppm higher than the present day.


Overview

The element
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent In chemistry, the valence (US spelling) or valency (British spelling) of an element is the measure of its combining capacity with o ...
plays a central role in climate and life on Earth. It is capable of moving among and between the
geosphere There are several conflicting usages of geosphere, variously defined. It may be taken as the collective name for the lithosphere, the hydrosphere, the cryosphere, and the atmosphere. The different collectives of the geosphere are able to exchan ...
,
cryosphere ] The cryosphere (from the Ancient Greek, Greek ''kryos'', "cold", "frost" or "ice" and ''sphaira'', "globe, ball") is an all-encompassing term for those portions of Earth's surface where water is in solid form, including sea ice, lake ice, ri ...
,
atmosphere An atmosphere () is a layer of gas or layers of gases that envelop a planet, and is held in place by the gravity of the planetary body. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A s ...
,
biosphere The biosphere (from Greek βίος ''bíos'' "life" and σφαῖρα ''sphaira'' "sphere"), also known as the ecosphere (from Greek οἶκος ''oîkos'' "environment" and σφαῖρα), is the worldwide sum of all ecosystems. It can also be ...
and
hydrosphere The hydrosphere () is the combined mass of water found on, under, and above the surface of a planet, minor planet, or natural satellite. Although Earth's hydrosphere has been around for about 4 billion years, it continues to change in shape. This ...
. This flow of carbon is referred to as the Earth's
carbon cycle The carbon cycle is the biogeochemical cycle by which carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and Earth's atmosphere, atmosphere of the Earth. Carbon is the main component of biological compounds as well as ...
. It is also intimately linked to the cycling of other elements and compounds. The ocean plays a fundamental role in Earth's carbon cycle, helping to regulate atmospheric CO2 concentration. The biological pump is a set of processes that transfer
organic carbon Total organic carbon (TOC) is the amount of carbon found in an organic compound and is often used as a non-specific indicator of water quality or cleanliness of pharmaceutical manufacturing equipment. TOC may also refer to the amount of organic c ...
from the surface to the deep ocean, and is at the heart of the
ocean carbon cycle The oceanic carbon cycle (or marine carbon cycle) is composed of processes that exchange carbon between various pools within the ocean as well as between the atmosphere, Earth interior, and the seafloor. The carbon cycle is a result of many inter ...
. The biological pump depends on the fraction of primary produced
organic matter Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
that survives degradation in the
euphotic zone The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological proc ...
and that is exported from surface water to the ocean interior, where it is mineralized to
inorganic carbon Total inorganic carbon (''C''T or TIC) is the sum of the inorganic carbon species. Carbon Chemical compound, compounds can be distinguished as either Organic compound, organic or inorganic, and Dissolved organic carbon, dissolved or Particulate ...
, with the result that carbon is transported against the gradient of
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
(DIC) from the surface to the deep ocean. This transfer occurs through physical mixing and transport of dissolved and
particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(POC), vertical migrations of organisms (
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
,
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
) and through gravitational settling of particulate organic carbon. Modified text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
The biological pump can be divided into three distinct phases, the first of which is the production of fixed carbon by planktonic
phototrophs Phototrophs () are organisms that carry out photon capture to produce complex organic compounds (e.g. carbohydrates) and acquire energy. They use the energy from light to carry out various cellular metabolic processes. It is a common misconcep ...
in the
euphotic The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological proc ...
(sunlit) surface region of the ocean. In these surface waters,
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...
use
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
(CO2),
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
(N),
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
(P), and other trace elements (
barium Barium is a chemical element with the symbol Ba and atomic number 56. It is the fifth element in group 2 and is a soft, silvery alkaline earth metal. Because of its high chemical reactivity, barium is never found in nature as a free element. Th ...
,
iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in f ...
,
zinc Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodi ...
, etc.) during photosynthesis to make
carbohydrates In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or may ...
,
lipids Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
, and
proteins Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
. Some plankton, (e.g.
coccolithophores Coccolithophores, or coccolithophorids, are single celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdo ...
and
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
) combine calcium (Ca) and dissolved carbonates ( carbonic acid and
bicarbonate In inorganic chemistry, bicarbonate (IUPAC-recommended nomenclature: hydrogencarbonate) is an intermediate form in the deprotonation of carbonic acid. It is a polyatomic anion with the chemical formula . Bicarbonate serves a crucial biochemic ...
) to form a calcium carbonate (CaCO3) protective coating. Once this carbon is fixed into soft or hard tissue, the organisms either stay in the euphotic zone to be recycled as part of the regenerative
nutrient cycle A nutrient cycle (or ecological recycling) is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cycli ...
or once they die, continue to the second phase of the biological pump and begin to sink to the ocean floor. The sinking particles will often form aggregates as they sink, greatly increasing the sinking rate. It is this aggregation that gives particles a better chance of escaping predation and decomposition in the water column and eventually making it to the sea floor. The fixed carbon that is decomposed by bacteria either on the way down or once on the sea floor then enters the final phase of the pump and is remineralized to be used again in
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through c ...
. The particles that escape these processes entirely are sequestered in the sediment and may remain there for millions of years. It is this sequestered carbon that is responsible for ultimately lowering atmospheric CO2. The diagram immediately above illustrates the components of the biological pump. Biology, physics and gravity interact to pump organic carbon into the deep sea. The processes of fixation of inorganic carbon in organic matter during photosynthesis, its transformation by food web processes (trophodynamics), physical mixing, transport and gravitational settling are referred to collectively as the biological pump. The biological pump is responsible for transforming
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
(DIC) into organic biomass and pumping it in
particulate Particulates – also known as atmospheric aerosol particles, atmospheric particulate matter, particulate matter (PM) or suspended particulate matter (SPM) – are microscopic particles of solid or liquid matter suspended in the air. The ter ...
or dissolved form into the deep ocean. Inorganic nutrients and carbon dioxide are fixed during photosynthesis by phytoplankton, which both release
dissolved organic matter Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOM) and are consumed by herbivorous zooplankton. Larger zooplankton - such as
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s -
egest Defecation (or defaecation) follows digestion, and is a necessary process by which organisms eliminate a solid, semisolid, or liquid waste material known as feces from the digestive tract via the anus. The act has a variety of names ranging fro ...
fecal pellet Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
s which can be reingested and sink or collect with other organic detritus into larger, more-rapidly-sinking aggregates. DOM is partially consumed by bacteria (black dots) and respired; the remaining
refractory DOM Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
is
advected In the field of physics, engineering, and earth sciences, advection is the transport of a substance or quantity by bulk motion of a fluid. The properties of that substance are carried with it. Generally the majority of the advected substance is al ...
and mixed into the deep sea. DOM and aggregates exported into the deep water are consumed and respired, thus returning organic carbon into the enormous deep ocean reservoir of DIC. About 1% of the particles leaving the surface ocean reach the seabed and are consumed, respired, or buried in the sediments. There, carbon is stored for millions of years. The net effect of these processes is to remove carbon in organic form from the surface and return it to DIC at greater depths, maintaining the surface-to-deep ocean gradient of DIC.
Thermohaline circulation Thermohaline circulation (THC) is a part of the large-scale ocean circulation that is driven by global density gradients created by surface heat and freshwater fluxes. The adjective ''thermohaline'' derives from '' thermo-'' referring to temper ...
returns deep-ocean DIC to the atmosphere on millennial timescales.


Primary production

The first step in the biological pump is the synthesis of both organic and inorganic carbon compounds by phytoplankton in the uppermost, sunlit layers of the ocean. Organic compounds in the form of sugars, carbohydrates, lipids, and proteins are synthesized during the process of
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
: CO2 + H2O + light → CH2O + O2 In addition to carbon, organic matter found in phytoplankton is composed of nitrogen, phosphorus and various
trace metals Trace metals are the metals subset of trace elements; that is, metals normally present in small but measurable amounts in animal and plant cells and tissues and that are a necessary part of nutrition and physiology. Many biometals are trace me ...
. The ratio of carbon to nitrogen and phosphorus varies from place to place, but has an average ratio near 106C:16N:1P, known as the
Redfield ratio The Redfield ratio or Redfield stoichiometry is the consistent atomic ratio of carbon, nitrogen and phosphorus found in marine phytoplankton and throughout the deep oceans. The term is named for American oceanographer Alfred C. Redfield who in 19 ...
. Trace metals such as magnesium, cadmium, iron, calcium, barium and copper are orders of magnitude less prevalent in phytoplankton organic material, but necessary for certain metabolic processes and therefore can be limiting nutrients in photosynthesis due to their lower abundance in the water column. Oceanic primary production accounts for about half of the carbon fixation carried out on Earth. Approximately 50–60 Pg of carbon are fixed by marine phytoplankton each year despite the fact that they comprise less than 1% of the total photosynthetic biomass on Earth. The majority of this carbon fixation (~80%) is carried out in the open ocean while the remaining amount occurs in the very productive
upwelling Upwelling is an oceanographic phenomenon that involves wind-driven motion of dense, cooler, and usually nutrient-rich water from deep water towards the ocean surface. It replaces the warmer and usually nutrient-depleted surface water. The nutr ...
regions of the ocean. Despite these productive regions producing 2 to 3 times as much fixed carbon per area, the open ocean accounts for greater than 90% of the ocean area and therefore is the larger contributor.


Forms of carbon


Dissolved and particulate carbon

Phytoplankton supports all life in the ocean as it converts inorganic compounds into organic constituents. This autotrophically produced biomass presents the foundation of the marine food web. In the diagram immediately below, the arrows indicate the various production (arrowhead pointing toward DOM pool) and removal processes of DOM (arrowhead pointing away), while the dashed arrows represent dominant biological processes involved in the transfer of DOM. Due to these processes, the fraction of labile DOM decreases rapidly with depth, whereas the refractory character of the DOM pool considerably increases during its export to the deep ocean. DOM, dissolved organic matter.Carlson CA (2002) ooks.google.com/books?id=D6TMKZOgldAC&q=%22Production+and+removal+processes%22&pg=PA91 "Production and removal processes" In: Hansell DA, Carlson CA (eds) ''Biogeochemistry of marine dissolved organic matter''. Academic Press, San Diego, pages 91–151. .


Ocean carbon pools

The marine biological pump depends on a number of key pools, components and processes that influence its functioning. There are four main pools of carbon in the ocean. Modified material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
*
Dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
(DIC) is the largest pool. It constitutes around 38,000 Pg C and includes dissolved carbon dioxide (CO2),
bicarbonate In inorganic chemistry, bicarbonate (IUPAC-recommended nomenclature: hydrogencarbonate) is an intermediate form in the deprotonation of carbonic acid. It is a polyatomic anion with the chemical formula . Bicarbonate serves a crucial biochemic ...
(),
carbonate A carbonate is a salt of carbonic acid (H2CO3), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word ''carbonate'' may also refer to a carbonate ester, an organic compound containing the carbonate g ...
(), and carbonic acid (). The equilibrium between carbonic acid and carbonate determines the pH of the seawater. Carbon dioxide dissolves easily in water and its solubility is inversely related to temperature. Dissolved CO2 is taken up in the process of photosynthesis, and can reduce the partial pressure of CO2 in the seawater, favouring drawdown from the atmosphere. The reverse process respiration, releases CO2 back into the water, can increase partial pressure of CO2 in the seawater, favouring release back to the atmosphere. The formation of calcium carbonate by organisms such as
coccolithophore Coccolithophores, or coccolithophorids, are single celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdo ...
s has the effect of releasing CO2 into the water. *
Dissolved organic carbon Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOC) is the next largest pool at around 662 Pg C. DOC can be classified according to its reactivity as refractory, semi-labile or labile. The labile pool constitutes around 0.2 Pg C, is bioavailable, and has a high production rate (~ 15−25 Pg C y−1). The refractory component is the biggest pool (~642 Pg C ± 32; but has a very low turnover rate (0.043 Pg C y−1). The turnover time for
refractory DOC Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
is thought to be greater than 1000 years. *
Particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(POC) constitutes around 2.3 Pg C, and is relatively small compared with DIC and DOC. Though small in size, this pool is highly dynamic, having the highest turnover rate of any organic carbon pool on the planet. Driven by
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through c ...
, it produces around 50 Pg C y−1 globally. It can be separated into living (e.g.
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...
,
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
,
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
) and non-living (e.g.
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
) material. Of these, the phytoplankton carbon is particularly important, because of its role in
marine primary production Marine primary production is the chemical synthesis in the ocean of organic compounds from atmospheric or dissolved carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it al ...
, and also because it serves as the food resource for all the larger organisms in the pelagic ecosystem. *
Particulate inorganic carbon Particulate inorganic carbon (PIC) can be contrasted with dissolved inorganic carbon (DIC), the other form of inorganic carbon found in the ocean. These distinctions are important in chemical oceanography. Particulate inorganic carbon is sometim ...
(PIC) is the smallest of the pools at around 0.03 Pg C. It is present in the form of calcium carbonate (CaCO3) in particulate form, and impacts the carbonate system and pH of the seawater. Estimates for PIC production are in the region of 0.8–1.4 Pg C y−1, with at least 65% of it being dissolved in the upper
water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...
, the rest contributing to deep sediments. Coccolithophores and
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
are estimated to be the dominant sources of PIC in the
open ocean The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
. The PIC pool is of particular importance due to its role in the ocean carbonate system, and in facilitating the export of carbon to the deep ocean through the
carbonate pump The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sediments.Sigman DM & GH ...
, whereby PIC is exported out of the
photic zone The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological proc ...
and deposited in the bottom sediments.


Calcium carbonate

Particulate inorganic carbon Particulate inorganic carbon (PIC) can be contrasted with dissolved inorganic carbon (DIC), the other form of inorganic carbon found in the ocean. These distinctions are important in chemical oceanography. Particulate inorganic carbon is sometim ...
(PIC) usually takes the form of calcium carbonate (CaCO3), and plays a key part in the ocean carbon cycle. This biologically fixed carbon is used as a protective coating for many planktonic species (coccolithophores, foraminifera) as well as larger marine organisms (mollusk shells). Calcium carbonate is also excreted at high rates during
osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
by fish, and can form in
whiting event A whiting event is a phenomenon that occurs when a suspended cloud of fine-grained calcium carbonate precipitates in water bodies, typically during summer months, as a result of photosynthetic microbiological activity or sediment disturbance. Th ...
s. While this form of carbon is not directly taken from the atmospheric budget, it is formed from dissolved forms of carbonate which are in equilibrium with CO2 and then responsible for removing this carbon via sequestration. CO2 + H2O → H2CO3 → H+ + HCO3 Ca2+ + 2HCO3 → CaCO3 + CO2 + H2O While this process does manage to fix a large amount of carbon, two units of
alkalinity Alkalinity (from ar, القلوي, al-qaly, lit=ashes of the saltwort) is the capacity of water to resist acidification. It should not be confused with basicity, which is an absolute measurement on the pH scale. Alkalinity is the strength of ...
are sequestered for every unit of sequestered carbon. The formation and sinking of CaCO3 therefore drives a surface to deep
alkalinity Alkalinity (from ar, القلوي, al-qaly, lit=ashes of the saltwort) is the capacity of water to resist acidification. It should not be confused with basicity, which is an absolute measurement on the pH scale. Alkalinity is the strength of ...
gradient which serves to raise the pH of surface waters, shifting the speciation of dissolved carbon to raise the
partial pressure In a mixture of gases, each constituent gas has a partial pressure which is the notional pressure of that constituent gas as if it alone occupied the entire volume of the original mixture at the same temperature. The total pressure of an ideal gas ...
of dissolved CO2 in surface waters, which actually raises atmospheric levels. In addition, the burial of CaCO3 in sediments serves to lower overall oceanic
alkalinity Alkalinity (from ar, القلوي, al-qaly, lit=ashes of the saltwort) is the capacity of water to resist acidification. It should not be confused with basicity, which is an absolute measurement on the pH scale. Alkalinity is the strength of ...
, tending to raise pH and thereby atmospheric CO2 levels if not counterbalanced by the new input of alkalinity from weathering. The portion of carbon that is permanently buried at the sea floor becomes part of the geologic record. Calcium carbonate often forms remarkable deposits that can then be raised onto land through tectonic motion as in the case with the
White Cliffs of Dover The White Cliffs of Dover is the region of English coastline facing the Strait of Dover and France. The cliff face, which reaches a height of , owes its striking appearance to its composition of chalk accented by streaks of black flint, deposi ...
in Southern England. These cliffs are made almost entirely of the plates of buried
coccolithophore Coccolithophores, or coccolithophorids, are single celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdo ...
s.Webb, Paul (2019) ''Introduction to Oceanography''
Chapter 12: Ocean Sediments
page 273–297, Rebus Community. Updated 2020.


Oceanic carbon cycle

Three main processes (or pumps) that make up the marine carbon cycle bring atmospheric
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
(CO2) into the ocean interior and distribute it through the oceans. These three pumps are: (1) the solubility pump, (2) the carbonate pump, and (3) the biological pump. The total active pool of carbon at the Earth's surface for durations of less than 10,000 years is roughly 40,000 gigatons C (Gt C, a
gigaton ''Gigaton'' is the eleventh studio album by American rock band Pearl Jam, released March 27, 2020. It was preceded by the singles "Dance of the Clairvoyants", " Superblood Wolfmoon" and "Quick Escape". It is the band's first studio album in six ...
is one billion tons, or the weight of approximately 6 million
blue whale The blue whale (''Balaenoptera musculus'') is a marine mammal and a baleen whale. Reaching a maximum confirmed length of and weighing up to , it is the largest animal known to have ever existed. The blue whale's long and slender body can ...
s), and about 95% (~38,000 Gt C) is stored in the ocean, mostly as
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
. The speciation of dissolved inorganic carbon in the marine carbon cycle is a primary controller of acid-base chemistry in the oceans.


Solubility pump

The biological pump is accompanied by a physico-chemical counterpart known as the
solubility pump In oceanic biogeochemistry, the solubility pump is a physico-chemical process that transports carbon as dissolved inorganic carbon (DIC) from the ocean's surface to its interior. Overview The solubility pump is driven by the coincidence of two ...
. This pump transports significant amounts of carbon in the form of
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
(DIC) from the ocean's surface to its interior. It involves physical and chemical processes only, and does not involve biological processes. The solubility pump is driven by the coincidence of two processes in the ocean: * The
solubility In chemistry, solubility is the ability of a substance, the solute, to form a solution with another substance, the solvent. Insolubility is the opposite property, the inability of the solute to form such a solution. The extent of the solubil ...
of
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
is a strong inverse function of seawater temperature (i.e. solubility is greater in cooler water) * The
thermohaline circulation Thermohaline circulation (THC) is a part of the large-scale ocean circulation that is driven by global density gradients created by surface heat and freshwater fluxes. The adjective ''thermohaline'' derives from '' thermo-'' referring to temper ...
is driven by the formation of deep water at high latitudes where seawater is usually cooler and denser Since deep water (that is, seawater in the ocean's interior) is formed under the same surface conditions that promote carbon dioxide solubility, it contains a higher concentration of dissolved inorganic carbon than might be expected from average surface concentrations. Consequently, these two processes act together to pump carbon from the atmosphere into the ocean's interior. One consequence of this is that when deep water upwells in warmer, equatorial latitudes, it strongly outgasses carbon dioxide to the atmosphere because of the reduced solubility of the gas.


Carbonate pump

The carbonate pump is sometimes referred to as the “hard tissue” component of the biological pump. Some surface marine organisms, like
coccolithophore Coccolithophores, or coccolithophorids, are single celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdo ...
s, produce hard structures out of calcium carbonate, a form of particulate inorganic carbon, by fixing bicarbonate. This fixation of DIC is an important part of the oceanic carbon cycle. Ca2+ + 2 HCO3 → CaCO3 + CO2 + H2O While the biological carbon pump fixes inorganic carbon (CO2) into
particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
in the form of sugar (C6H12O6), the carbonate pump fixes inorganic bicarbonate and causes a net release of CO2. In this way, the carbonate pump could be termed the carbonate counter pump. It works counter to the biological pump by counteracting the CO2 flux into the biological pump.


Continental shelf pump

The
continental shelf pump In oceanic biogeochemistry, the continental shelf pump is proposed to operate in the shallow waters of the continental shelf, continental shelves, acting as a mechanism to transport carbon (as either dissolved or particulate material) from surfac ...
is proposed as operating in the shallow waters of the
continental shelves A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
as a mechanism transporting carbon (dissolved or particulate) from the continental waters to the interior of the adjacent deep ocean. As originally formulated, the pump is thought to occur where the
solubility pump In oceanic biogeochemistry, the solubility pump is a physico-chemical process that transports carbon as dissolved inorganic carbon (DIC) from the ocean's surface to its interior. Overview The solubility pump is driven by the coincidence of two ...
interacts with cooler, and therefore denser water from the shelf floor which feeds down the
continental slope A continental margin is the outer edge of continental crust abutting oceanic crust under coastal waters. It is one of the three major zones of the ocean floor, the other two being deep-ocean basins and mid-ocean ridges. The continental margin ...
into the neighbouring deep ocean. The shallowness of the continental shelf restricts the
convection Convection is single or multiphase fluid flow that occurs spontaneously due to the combined effects of material property heterogeneity and body forces on a fluid, most commonly density and gravity (see buoyancy). When the cause of the convec ...
of cooling water, so the cooling can be greater for continental shelf waters than for neighbouring open ocean waters. These cooler waters promote the
solubility pump In oceanic biogeochemistry, the solubility pump is a physico-chemical process that transports carbon as dissolved inorganic carbon (DIC) from the ocean's surface to its interior. Overview The solubility pump is driven by the coincidence of two ...
and lead to an increased storage of
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
. This extra carbon storage is further augmented by the increased biological production characteristic of shelves.Wollast, R. (1998). Evaluation and comparison of the global carbon cycle in the coastal zone and in the open ocean, p. 213-252. In K. H. Brink and A. R. Robinson (eds.), ''The Global Coastal Ocean''. John Wiley & Sons. The dense, carbon-rich shelf waters then sink to the shelf floor and enter the sub-surface layer of the open ocean via
isopycnal Isopycnals are layers within the ocean that are stratified based on their densities and can be shown as a line connecting points of a specific density or potential density on a graph. Isopycnals are often displayed graphically to help visualize " ...
mixing. As the sea level rises in response to global warming, the surface area of the shelf seas will grow and in consequence the strength of the shelf sea pump should increase.


Processes in the biological pump

In the diagram on the right, phytoplankton convert CO2, which has dissolved from the atmosphere into the surface oceans (90 Gt yr−1), into
particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(POC) during
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through c ...
(~ 50 Gt C yr−1). Phytoplankton are then consumed by
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s,
krill Krill are small crustaceans of the order Euphausiacea, and are found in all the world's oceans. The name "krill" comes from the Norwegian Norwegian, Norwayan, or Norsk may refer to: *Something of, from, or related to Norway, a country in n ...
and other small zooplankton grazers, which in turn are preyed upon by higher
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...
s. Any unconsumed phytoplankton form aggregates, and along with zooplankton faecal pellets, sink rapidly and are exported out of the
mixed layer The oceanic or limnological mixed layer is a layer in which active turbulence has homogenized some range of depths. The surface mixed layer is a layer where this turbulence is generated by winds, surface heat fluxes, or processes such as evaporat ...
(< 12 Gt C yr−1 14). Krill, copepods, zooplankton and microbes intercept phytoplankton in the surface ocean and sinking detrital particles at depth, consuming and respiring this POC to CO2 (
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
, DIC), such that only a small proportion of surface-produced carbon sinks to the deep ocean (i.e., depths > 1000 m). As krill and smaller zooplankton feed, they also physically fragment particles into small, slower- or non-sinking pieces (via sloppy feeding, coprorhexy if fragmenting faeces), retarding POC export. This releases
dissolved organic carbon Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOC) either directly from cells or indirectly via bacterial solubilisation (yellow circle around DOC). Bacteria can then
remineralise In biogeochemistry, remineralisation (or remineralization) refers to the breakdown or transformation of organic matter (those molecules derived from a biological source) into its simplest Inorganic compound, inorganic forms. These transformations ...
the DOC to DIC (CO2, microbial gardening).Cavan, E.L., Belcher, A., Atkinson, A., Hill, S.L., Kawaguchi, S., McCormack, S., Meyer, B., Nicol, S., Ratnarajah, L., Schmidt, K. and Steinberg, D.K. (2019) "The importance of Antarctic krill in biogeochemical cycles". ''Nature communications'', 10(1): 1–13. . Modified text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
The biological carbon pump is one of the chief determinants of the vertical distribution of carbon in the oceans and therefore of the surface partial pressure of CO2 governing air-sea CO2 exchange. It comprises phytoplankton cells, their consumers and the bacteria that assimilate their waste and plays a central role in the global carbon cycle by delivering carbon from the atmosphere to the deep sea, where it is concentrated and sequestered for centuries. Photosynthesis by phytoplankton lowers the partial pressure of CO2 in the upper ocean, thereby facilitating the absorption of CO2 from the atmosphere by generating a steeper CO2 gradient. It also results in the formation of
particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(POC) in the euphotic layer of the
epipelagic zone The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological proc ...
(0–200 m depth). The POC is processed by microbes, zooplankton and their consumers into fecal pellets, organic aggregates (“marine snow”) and other forms, which are thereafter exported to the
mesopelagic The mesopelagic zone (Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins ...
(200–1000 m depth) and
bathypelagic zone The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above, and the abyssopelagic below. The bathypelagic ...
s by sinking and vertical migration by zooplankton and fish. Although primary production includes both dissolved and
particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(DOC and POC respectively), only POC leads to efficient carbon export to the ocean interior, whereas the DOC fraction in surface waters is mostly recycled by bacteria. However, a more biologically resistant DOC fraction produced in the euphotic zone (accounting for 15–20% of net community productivity), is not immediately mineralized by microbes and accumulates in the ocean surface as biologically
semi-labile DOC Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
. This semi-labile DOC undergoes net export to the deep ocean, thus constituting a dynamic part of the biological carbon pump. The efficiency of DOC production and export varies across oceanographic regions, being more prominent in the
oligotrophic An oligotroph is an organism that can live in an environment that offers very low levels of nutrients. They may be contrasted with copiotrophs, which prefer nutritionally rich environments. Oligotrophs are characterized by slow growth, low rates of ...
subtropical oceans. The overall efficiency of the biological carbon pump is mostly controlled by the export of POC.


Marine snow

Most carbon incorporated in organic and inorganic biological matter is formed at the sea surface where it can then start sinking to the ocean floor. The deep ocean gets most of its nutrients from the higher water column when they sink down in the form of marine snow. This is made up of dead or dying animals and microbes, fecal matter, sand and other inorganic material. A single phytoplankton cell has a sinking rate around one metre per day. Given that the average depth of the ocean is about four kilometres, it can take over ten years for these cells to reach the ocean floor. However, through processes such as coagulation and expulsion in predator fecal pellets, these cells form aggregates. These aggregates, known as
marine snow In the deep ocean, marine snow (also known as "ocean dandruff") is a continuous shower of mostly organic detritus falling from the upper layers of the water column. It is a significant means of exporting energy from the light-rich photic zone to ...
, have sinking rates orders of magnitude greater than individual cells and complete their journey to the deep in a matter of days. In the diagram on the right,
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...
fix CO2 in the
euphotic The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological proc ...
zone using solar energy and produce
particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(POC). POC formed in the euphotic zone is processed by microbes, zooplankton and their consumers into organic aggregates (marine snow), which is thereafter exported to the
mesopelagic The mesopelagic zone (Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins ...
(200–1000 m depth) and
bathypelagic zone The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above, and the abyssopelagic below. The bathypelagic ...
s by sinking and
vertical migration Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The word ''diel'' comes from the Latin ''dies'' day, and means a 24- ...
by zooplankton and fish. Export flux is defined as the
sedimentation Sedimentation is the deposition of sediments. It takes place when particles in suspension settle out of the fluid in which they are entrained and come to rest against a barrier. This is due to their motion through the fluid in response to the ...
out of the surface layer (at approximately 100 m depth) and sequestration flux is the sedimentation out of the mesopelagic zone (at approximately 1000 m depth). A portion of the POC is respired back to CO2 in the oceanic
water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...
at depth, mostly by
heterotrophic A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
microbes and zooplankton, thus maintaining a vertical gradient in concentration of
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
(DIC). This deep-ocean DIC returns to the atmosphere on millennial timescales through
thermohaline circulation Thermohaline circulation (THC) is a part of the large-scale ocean circulation that is driven by global density gradients created by surface heat and freshwater fluxes. The adjective ''thermohaline'' derives from '' thermo-'' referring to temper ...
. Between 1% and 40% of the primary production is exported out of the euphotic zone, which attenuates exponentially towards the base of the mesopelagic zone and only about 1% of the surface production reaches the sea floor. Of the 50–60 Pg of carbon fixed annually, roughly 10% leaves the surface mixed layer of the oceans, while less than 0.5% of eventually reaches the sea floor. Most is retained in regenerated production in the euphotic zone and a significant portion is remineralized in midwater processes during particle sinking. The portion of carbon that leaves the surface mixed layer of the ocean is sometimes considered "sequestered", and essentially removed from contact with the atmosphere for many centuries. However, work also finds that, in regions such as the
Southern Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-small ...
, much of this carbon can quickly (within decades) come back into contact with the atmosphere. Budget calculations of the biological carbon pump are based on the ratio between
sedimentation Sedimentation is the deposition of sediments. It takes place when particles in suspension settle out of the fluid in which they are entrained and come to rest against a barrier. This is due to their motion through the fluid in response to the ...
(carbon export) and
remineralization In biogeochemistry, remineralisation (or remineralization) refers to the breakdown or transformation of organic matter (those molecules derived from a biological source) into its simplest Inorganic compound, inorganic forms. These transformations ...
(release to the atmosphere). Modified text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
It has been estimated that sinking particles export up to 25% of the carbon captured by phytoplankton in the surface ocean to deeper water layers. About 20% of this export (5% of surface values) is buried in the ocean sediments mainly due to their mineral ballast. During the sinking process, these organic particles are hotspots of microbial activity and represent important loci for organic matter mineralization and nutrient redistribution in the water column.


Biomineralization


Ballast minerals

Observations have shown that fluxes of ballast minerals (calcium carbonate, opal, and lithogenic material) and organic carbon fluxes are closely correlated in the
bathypelagic zone The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above, and the abyssopelagic below. The bathypelagic ...
s of the ocean. A large fraction of particulate organic matter occurs in the form of marine snow aggregates (>0.5 mm) composed of phytoplankton, detritus, inorganic mineral grains, and fecal pellets in the ocean. Formation and sinking of these aggregates drive the biological carbon pump via export and sedimentation of organic matter from the surface mixed layer to the deep ocean and sediments. The fraction of organic matter that leaves the upper mixed layer of the ocean is, among other factors, determined by the sinking velocity and microbial
remineralisation In biogeochemistry, remineralisation (or remineralization) refers to the breakdown or transformation of organic matter (those molecules derived from a biological source) into its simplest inorganic forms. These transformations form a crucial link ...
rate of these aggregates. Recent observations have shown that the fluxes of ballast minerals (calcium carbonate, opal, and lithogenic material) and the organic carbon fluxes are closely correlated in the bathypelagic zones of the ocean. This has led to the hypothesis that organic carbon export is determined by the presence of ballast minerals within settling aggregates. Modified text was copied from this source, which is available under
Creative Commons Attribution 3.0 International License
Mineral ballasting is associated with about 60% of the flux of particulate organic carbon (POC) in the high-latitude North Atlantic, and with about 40% of the flux in the Southern Ocean. Strong correlations exist also in the deep ocean between the presence of ballast minerals and the flux of POC. This suggests ballast minerals enhance POC flux by increasing the sink rate of ballasted aggregates. Ballast minerals could additionally provide aggregated organic matter some protection from degradation. It has been proposed that organic carbon is better preserved in sinking particles due to increased aggregate density and sinking velocity when ballast minerals are present and/or via protection of the organic matter due to quantitative association to ballast minerals. In 2002, Klaas and Archer observed that about 83% of the global particulate organic carbon (POC) fluxes were associated with
carbonate A carbonate is a salt of carbonic acid (H2CO3), characterized by the presence of the carbonate ion, a polyatomic ion with the formula . The word ''carbonate'' may also refer to a carbonate ester, an organic compound containing the carbonate g ...
, and suggested carbonate was a more efficient ballast mineral as compared to opal and terrigenous material. They hypothesized that the higher density of calcium carbonate compared to that of opal and the higher abundance of calcium carbonate relative to terrigenous material might be the reason for the efficient ballasting by calcium carbonate. However, the direct effects of ballast minerals on sinking velocity and degradation rates in sinking aggregates are still unclear. A 2008 study demonstrated copepod fecal pellets produced on a diet of diatoms or coccolithophorids show higher sinking velocities as compared to pellets produced on a nanoflagellate diet. Carbon-specific respiration rates in pellets, however, were similar and independent of mineral content. These results suggest differences in mineral composition do not lead to differential protection of POC against microbial degradation, but the enhanced sinking velocities may result in up to 10-fold higher carbon preservation in pellets containing
biogenic mineral Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, often to harden or stiffen existing tissues. Such tissues are called mineralized tissues. It is an extremely widespread phenomenon; ...
s as compared to that of pellets without biogenic minerals Minerals seem to enhance the
flocculation Flocculation, in the field of chemistry, is a process by which colloidal particles come out of suspension to sediment under the form of floc or flake, either spontaneously or due to the addition of a clarifying agent. The action differs from pr ...
of phytoplankton aggregates and may even act as a catalyst in aggregate formation. However, it has also been shown that incorporation of minerals can cause aggregates to fragment into smaller and denser aggregates. This can potentially lower the sinking velocity of the aggregated organic material due to the reduced aggregate sizes, and, thus, lower the total export of organic matter. Conversely, if the incorporation of minerals increases the aggregate density, its size-specific sinking velocity may also increase, which could potentially increase the carbon export. Therefore, there is still a need for better quantitative investigations of how the interactions between minerals and organic aggregates affect the degradation and sinking velocity of the aggregates and, hence, carbon sequestration in the ocean.


Remineralisation

Remineralisation In biogeochemistry, remineralisation (or remineralization) refers to the breakdown or transformation of organic matter (those molecules derived from a biological source) into its simplest inorganic forms. These transformations form a crucial link ...
refers to the breakdown or transformation of
organic matter Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
(those molecules derived from a biological source) into its simplest
inorganic In chemistry, an inorganic compound is typically a chemical compound that lacks carbon–hydrogen bonds, that is, a compound that is not an organic compound. The study of inorganic compounds is a subfield of chemistry known as '' inorganic chemist ...
forms. These transformations form a crucial link within
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s as they are responsible for liberating the energy stored in
organic molecules In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen or carbon-carbon bonds. Due to carbon's ability to catenate (form chains with other carbon atoms), millions of organic compounds are known. The s ...
and recycling matter within the system to be reused as
nutrient A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excret ...
s by other
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
s. What fraction does escape remineralisation varies depending on the location. For example, in the North Sea, values of carbon deposition are ~1% of primary production while that value is <0.5% in the open oceans on average. Therefore, most of nutrients remain in the water column, recycled by the biota.
Heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic organisms will utilize the materials produced by the
autotroph An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...
ic (and
chemotroph A Chemotroph is an organism that obtains energy by the oxidation of electron donors in their environments. These molecules can be organic ( chemoorganotrophs) or inorganic (chemolithotrophs). The chemotroph designation is in contrast to phototro ...
ic) organisms and via respiration will remineralise the compounds from the organic form back to inorganic, making them available for primary producers again. For most areas of the ocean, the highest rates of carbon remineralisation occur at depths between in the water column, decreasing down to about 1,200 m where remineralisation rates remain pretty constant at 0.1 μmol kg−1 yr−1. This provides the most nutrients available for primary producers within the photic zone, though it leaves the upper surface waters starved of inorganic nutrients. Most remineralisation is done with
dissolved organic carbon Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOC). Studies have shown that it is larger sinking particles that transport matter down to the sea floor while suspended particles and dissolved organics are mostly consumed by remineralisation. This happens in part due to the fact that organisms must typically ingest nutrients smaller than they are, often by orders of magnitude. With the microbial community making up 90% of marine biomass, it is particles smaller than the microbes (on the order of ) that will be taken up for remineralisation.


Key role of phytoplankton

Marine phytoplankton perform half of all photosynthesis on Earth and directly influence global biogeochemical cycles and the climate, yet how they will respond to future global change is unknown. Carbon dioxide is one of the principal drivers of global change and has been identified as one of the major challenges in the 21st century. Carbon dioxide (CO2) generated during anthropogenic activities such as deforestation and burning of fossil fuels for energy generation rapidly dissolves in the surface ocean and lowers seawater pH, while CO2 remaining in the atmosphere increases global temperatures and leads to increased ocean thermal stratification. While CO2 concentration in the atmosphere is estimated to be about 270 ppm before the industrial revolution, it has currently increased to about 400 ppm and is expected to reach 800–1000 ppm by the end of this century according to the "business as usual" CO2 emission scenario. Marine ecosystems are a major sink for atmospheric CO2 and take up similar amount of CO2 as terrestrial ecosystems, currently accounting for the removal of nearly one third of anthropogenic CO2 emissions from the atmosphere. The net transfer of CO2 from the atmosphere to the oceans and then
sediments Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand a ...
, is mainly a direct consequence of the combined effect of the solubility and the biological pump. While the
solubility pump In oceanic biogeochemistry, the solubility pump is a physico-chemical process that transports carbon as dissolved inorganic carbon (DIC) from the ocean's surface to its interior. Overview The solubility pump is driven by the coincidence of two ...
serves to concentrate
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
(CO2 plus bicarbonate and carbonate ions) in the deep oceans, the biological carbon pump (a key natural process and a major component of the global carbon cycle that regulates atmospheric CO2 levels) transfers both organic and inorganic carbon fixed by
primary producers An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...
(phytoplankton) in the
euphotic zone The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological proc ...
to the ocean interior and subsequently to the underlying sediments. Thus, the biological pump takes carbon out of contact with the atmosphere for several thousand years or longer and maintains atmospheric CO2 at significantly lower levels than would be the case if it did not exist. An ocean without a biological pump, which transfers roughly 11 Gt C yr−1 into the ocean's interior, would result in atmospheric CO2 levels ~400 ppm higher than present day. Passow and Carlson defined sedimentation out of the surface layer (at approximately 100 m depth) as the "export flux" and that out of the
mesopelagic zone The mesopelagic zone (Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins ...
(at approximately 1000 m depth) as the "sequestration flux". Once carbon is transported below the mesopelagic zone, it remains in the deep sea for 100 years or longer, hence the term “sequestration” flux. According to the modelling results of Buesseler and Boyd between 1% and 40% of the primary production is exported out of the euphotic zone, which attenuates exponentially towards the base of the mesopelagic zone and only about 1% of the surface production reaches the sea floor. The export efficiency of
particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(POC) shows regional variability. For instance, in the North Atlantic, over 40% of net primary production is exported out of the euphotic zone as compared to only 10% in the South Pacific, and this is driven in part by the composition of the phytoplankton community including cell size and composition (see below). Exported organic carbon is remineralized, that is, respired back to CO2 in the oceanic water column at depth, mainly by heterotrophic microbes and zooplankton. Thus, the biological carbon pump maintains a vertical gradient in the concentration of
dissolved inorganic carbon Dissolved inorganic carbon (DIC) is the sum of the aqueous species of inorganic carbon in a solution. Carbon compounds can be distinguished as either organic or inorganic, and as dissolved or particulate, depending on their composition. Organic c ...
(DIC), with higher values at increased ocean depth. This deep-ocean DIC returns to the atmosphere on millennial timescales through
thermohaline circulation Thermohaline circulation (THC) is a part of the large-scale ocean circulation that is driven by global density gradients created by surface heat and freshwater fluxes. The adjective ''thermohaline'' derives from '' thermo-'' referring to temper ...
. Modified text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
In 2001, Hugh et al. expressed the efficiency of the biological pump as the amount of carbon exported from the surface layer (export production) divided by the total amount produced by photosynthesis (overall production). Modelling studies by Buesseler and Boyd revealed that the overall transfer efficiency of the biological pump is determined by a combination of factors: seasonality; the composition of phytoplankton species; the fragmentation of particles by zooplankton; and the solubilization of particles by microbes. In addition, the efficiency of the biological pump is also dependent on the aggregation and disaggregation of organic-rich aggregates and interaction between POC aggregates and suspended “ballast” minerals. Ballast minerals (silicate and carbonate biominerals and dust) are the major constituents of particles that leave the ocean surface via sinking. They are typically denser than seawater and most organic matter, thus, providing a large part of the density differential needed for sinking of the particles. Aggregation of particles increases vertical flux by transforming small suspended particles into larger, rapidly-sinking ones. It plays an important role in the sedimentation of phytodetritus from surface layer phytoplankton blooms. As illustrated by Turner in 2015, the vertical flux of sinking particles is mainly due to a combination of fecal pellets, marine snow and direct sedimentation of phytoplankton blooms, which are typically composed of diatoms, coccolithophorids, dinoflagellates and other plankton. Marine snow comprises macroscopic organic aggregates >500 µm in size and originates from clumps of aggregated phytoplankton (phytodetritus), discarded appendicularian houses, fecal matter and other miscellaneous detrital particles, Appendicularians secrete mucous feeding structures or “houses” to collect food particles and discard and renew them up to 40 times a day . Discarded appendicularian houses are highly abundant (thousands per m3 in surface waters) and are microbial hotspots with high concentrations of bacteria, ciliates, flagellates and phytoplankton. These discarded houses are therefore among the most important sources of aggregates directly produced by zooplankton in terms of carbon cycling potential. The composition of the phytoplankton community in the euphotic zone largely determines the quantity and quality of organic matter that sinks to depth. The main functional groups of marine phytoplankton that contribute to export production include nitrogen fixers (
diazotrophic Diazotrophs are bacteria and archaea that fix gaseous nitrogen in the atmosphere into a more usable form such as ammonia. A diazotroph is a microorganism that is able to grow without external sources of fixed nitrogen. Examples of organisms that d ...
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
), silicifiers (diatoms) and
calcifier Marine biogenic calcification is the process by which marine organisms such as oysters and clams form calcium carbonate. Seawater is full of dissolved compounds, ions and nutrients that organisms can use for energy and, in the case of calcificati ...
s (coccolithophores). Each of these phytoplankton groups differ in the size and composition of their cell walls and coverings, which influence their sinking velocities. For example, autotrophic picoplankton (0.2–2 µm in diameter)—which include taxa such as cyanobacteria (e.g., ''
Prochlorococcus ''Prochlorococcus'' is a genus of very small (0.6  μm) marine cyanobacteria with an unusual pigmentation ( chlorophyll ''a2'' and ''b2''). These bacteria belong to the photosynthetic picoplankton and are probably the most abundant photosynth ...
'' spp. and ''
Synechococcus ''Synechococcus'' (from the Greek ''synechos'', in succession, and the Greek ''kokkos'', granule) is a unicellular cyanobacterium that is very widespread in the marine environment. Its size varies from 0.8 to 1.5 µm. The photosynthetic c ...
'' spp.) and
prasinophyte The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapman ...
s (various genera of eukaryotes <2 µm)—are believed to contribute much less to carbon export from surface layers due to their small size, slow sinking velocities (<0.5 m/day) and rapid turnover in the microbial loop. In contrast, larger phytoplankton cells such as diatoms (2–500 µm in diameter) are very efficient in transporting carbon to depth by forming rapidly sinking aggregates. They are unique among phytoplankton, because they require Si in the form of silicic acid (Si(OH)4) for growth and production of their frustules, which are made of biogenic silica (bSiO2) and act as ballast. According to the reports of Miklasz and Denny, the sinking velocities of diatoms can range from 0.4 to 35 m/day. Analogously, coccolithophores are covered with calcium carbonate plates called 'coccoliths', which are central to aggregation and ballasting, producing sinking velocities of nearly 5 m/day. Although it has been assumed that
picophytoplankton Photosynthetic picoplankton or picophytoplankton is the fraction of the phytoplankton performing photosynthesis composed of cells between 0.2 and 2 µm in size ( picoplankton). It is especially important in the central oligotrophic regions of ...
, characterizing vast
oligotrophic An oligotroph is an organism that can live in an environment that offers very low levels of nutrients. They may be contrasted with copiotrophs, which prefer nutritionally rich environments. Oligotrophs are characterized by slow growth, low rates of ...
areas of the ocean, do not contribute substantially to the particulate organic carbon (POC) flux, in 2007 Richardson and Jackson suggested that all phytoplankton, including picoplankton cells, contribute equally to POC export. They proposed alternative pathways for picoplankton carbon cycling, which rely on aggregation as a mechanism for both direct sinking (the export of picoplankton as POC) and mesozooplankton- or large filter feeder-mediated sinking of picoplankton-based production.


Zooplankton grazing


Sloppy feeding

In addition to linking primary producers to higher trophic levels in marine food webs, zooplankton also play an important role as “recyclers” of carbon and other nutrients that significantly impact marine biogeochemical cycles, including the biological pump. This is particularly the case with
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s and
krill Krill are small crustaceans of the order Euphausiacea, and are found in all the world's oceans. The name "krill" comes from the Norwegian Norwegian, Norwayan, or Norsk may refer to: *Something of, from, or related to Norway, a country in n ...
, and is especially important in oligotrophic waters of the open ocean. Through sloppy feeding, excretion, egestion, and leaching of fecal pellets, zooplankton release dissolved organic matter (DOM) which controls DOM cycling and supports the microbial loop. Absorption efficiency, respiration, and prey size all further complicate how zooplankton are able to transform and deliver carbon to the deep ocean. Excretion and sloppy feeding (the physical breakdown of food source) make up 80% and 20% of crustacean zooplankton-mediated DOM release respectively. In the same study, fecal pellet leaching was found to be an insignificant contributor. For protozoan grazers, DOM is released primarily through excretion and egestion and gelatinous zooplankton can also release DOM through the production of mucus. Leaching of fecal pellets can extend from hours to days after initial egestion and its effects can vary depending on food concentration and quality. Various factors can affect how much DOM is released from zooplankton individuals or populations.


Fecal pellets

The fecal pellets of zooplankton can be important vehicles for the transfer of particulate organic carbon (POC) to the deep ocean, often making large contributions to the carbon sequestration. The size distribution of the copepod community indicates high numbers of small fecal pellets are produced in the
epipelagic The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological proc ...
. However, small fecal pellets are rare in the deeper layers, suggesting they are not transferred efficiently to depth. This means small fecal pellets make only minor contributions to fecal pellet fluxes in the meso- and bathypelagic, particularly in terms of carbon. In a study is focussed on the
Scotia Sea The Scotia Sea is a sea located at the northern edge of the Southern Ocean at its boundary with the South Atlantic Ocean. It is bounded on the west by the Drake Passage and on the north, east, and south by the Scotia Arc, an undersea ridge and i ...
, which contains some of the most productive regions in the Southern Ocean, the dominant fecal pellets in the upper
mesopelagic The mesopelagic zone (Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins ...
were cylindrical and elliptical, while
ovoid An oval () is a closed curve in a plane which resembles the outline of an egg. The term is not very specific, but in some areas (projective geometry, technical drawing, etc.) it is given a more precise definition, which may include either one or ...
fecal pellets were dominant in the
bathypelagic The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above, and the abyssopelagic below. The bathypelagic ...
. The change in fecal pellet morphology, as well as size distribution, points to the repacking of surface fecal pellets in the mesopelagic and in situ production in the lower meso- and bathypelagic, which may be augmented by inputs of fecal pellets via zooplankton vertical migrations. This suggests the flux of carbon to the deeper layers within the Southern Ocean is strongly modulated by meso- and bathypelagic zooplankton, meaning that the community structure in these zones has a major impact on the efficiency of the fecal pellet transfer to ocean depths.
Absorption efficiency Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
(AE) is the proportion of food absorbed by plankton that determines how available the consumed organic materials are in meeting the required physiological demands. Depending on the feeding rate and prey composition, variations in AE may lead to variations in fecal pellet production, and thus regulates how much organic material is recycled back to the marine environment. Low feeding rates typically lead to high AE and small, dense pellets, while high feeding rates typically lead to low AE and larger pellets with more organic content. Another contributing factor to DOM release is respiration rate. Physical factors such as oxygen availability, pH, and light conditions may affect overall oxygen consumption and how much carbon is loss from zooplankton in the form of respired CO2. The relative sizes of zooplankton and prey also mediate how much carbon is released via sloppy feeding. Smaller prey are ingested whole, whereas larger prey may be fed on more “sloppily”, that is more biomatter is released through inefficient consumption. There is also evidence that diet composition can impact nutrient release, with carnivorous diets releasing more
dissolved organic carbon Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOC) and ammonium than omnivorous diets.


Microbial loop


Bacterial lysis

The
microbial loop The microbial loop describes a trophic pathway where, in aquatic systems, dissolved organic carbon (DOC) is returned to higher trophic levels via its incorporation into bacterial biomass, and then coupled with the classic food chain formed by phy ...
describes a trophic pathway in the marine
microbial food web The microbial food web refers to the combined trophic interactions among microbes in aquatic environments. These microbes include viruses, bacteria, algae, heterotrophic protists (such as ciliates and flagellates).Mostajir B, Amblard C, Buffan-D ...
where
dissolved organic carbon Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOC) is returned to higher trophic levels via its incorporation into bacterial biomass, and then coupled with the classic food chain formed by
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...
-
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
-
nekton Nekton or necton (from the ) refers to the actively swimming aquatic organisms in a body of water. The term was proposed by German biologist Ernst Haeckel to differentiate between the active swimmers in a body of water, and the passive organisms ...
. The term microbial loop was coined by
Farooq Azam Farooq Azam (born in Lahore, Pakistan ) is a researcher in the field of marine microbiology. He is a Distinguished Professor at the Scripps Institution of Oceanography, at the University of California San Diego. Farooq Azam grew up in Lahore and ...
,
Tom Fenchel Tom or TOM may refer to: * Tom (given name), a diminutive of Thomas or Tomás or an independent Aramaic given name (and a list of people with the name) Characters * Tom Anderson, a character in '' Beavis and Butt-Head'' * Tom Beck, a character ...
et al. in 1983 to include the role played by bacteria in the carbon and nutrient cycles of the marine environment. In general, dissolved organic carbon is introduced into the ocean environment from
bacterial lysis Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
, the leakage or exudation of fixed carbon from phytoplankton (e.g., mucilaginous exopolymer from
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s), sudden cell senescence, sloppy feeding by zooplankton, the excretion of waste products by aquatic animals, or the breakdown or dissolution of organic particles from terrestrial plants and soils. Bacteria in the microbial loop decompose this particulate detritus to utilize this energy-rich matter for growth. Since more than 95% of organic matter in marine ecosystems consists of polymeric, high
molecular weight A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioch ...
(HMW) compounds (e.g., protein, polysaccharides, lipids), only a small portion of total
dissolved organic matter Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOM) is readily utilizable to most marine organisms at higher trophic levels. This means that dissolved organic carbon is not available directly to most marine organisms;
marine bacteria Marine prokaryotes are marine bacteria and marine archaea. They are defined by their habitat as prokaryotes that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. All cellular ...
introduce this organic carbon into the food web, resulting in additional energy becoming available to higher trophic levels.


Viral shunt

As much as 25% of the primary production from phytoplankton in the global oceans may be recycled within the microbial loop through
viral shunt The viral shunt is a mechanism that prevents marine microbial particulate organic matter (POM) from migrating up trophic levels by recycling them into dissolved organic matter (DOM), which can be readily taken up by microorganisms. The DOM rec ...
ing. The viral shunt is a mechanism whereby
marine viruses Marine viruses are defined by their habitat as viruses that are found in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Viruses are small infectious agents that can only replicate i ...
prevent microbial
particulate organic matter Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(POM) from migrating up
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...
s by recycling them into
dissolved organic matter Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOM), which can be readily taken up by microorganisms. The DOM recycled by the viral shunt pathway is comparable to the amount generated by the other main sources of marine DOM. Viruses can easily infect microorganisms in the microbial loop due to their relative abundance compared to microbes. Prokaryotic and eukaryotic mortality contribute to carbon nutrient recycling through
cell lysis Lysis ( ) is the breaking down of the membrane of a cell, often by viral, enzymic, or osmotic (that is, "lytic" ) mechanisms that compromise its integrity. A fluid containing the contents of lysed cells is called a ''lysate''. In molecular bio ...
. There is evidence as well of nitrogen (specifically ammonium) regeneration. This nutrient recycling helps stimulates microbial growth.


Macroorganisms


Jelly fall

Jelly-falls Jelly-falls are marine carbon cycling events whereby gelatinous zooplankton, primarily cnidarians, sink to the seafloor and enhance carbon and nitrogen fluxes via rapidly sinking particulate organic matter. These events provide nutrition to be ...
are marine
carbon cycling The carbon cycle is the biogeochemical cycle by which carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of the Earth. Carbon is the main component of biological compounds as well as a major component ...
events whereby
gelatinous zooplankton Gelatinous zooplankton are fragile animals that live in the water column in the ocean. Their delicate bodies have no hard parts and are easily damaged or destroyed. Gelatinous zooplankton are often transparent. All jellyfish are gelatinous zoopla ...
, primarily
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
ns, sink to the seafloor and enhance carbon and nitrogen fluxes via rapidly sinking
particulate organic matter Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
. These events provide nutrition to
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
megafauna In terrestrial zoology, the megafauna (from Greek μέγας ''megas'' "large" and New Latin ''fauna'' "animal life") comprises the large or giant animals of an area, habitat, or geological period, extinct and/or extant. The most common threshold ...
and
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
. Jelly-falls have been implicated as a major “gelatinous pathway” for the sequestration of
labile Lability refers to something that is constantly undergoing change or is likely to undergo change. Biochemistry In reference to biochemistry, this is an important concept as far as kinetics is concerned in metalloproteins. This can allow for th ...
biogenic carbon through the biological pump. These events are common in protected areas with high levels of primary production and water quality suitable to support cnidarian species. These areas include
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environment ...
and several studies have been conducted in
fjords of Norway This list of Norwegian fjords shows many of the fjords in Norway. In total, there are about 1,190 fjords in Norway and the Svalbard islands. The sortable list includes the lengths and locations of those fjords. Fjords See also * List of gla ...
.


Whale pump

Whale Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
s and other
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their reli ...
s also enhance primary productivity in their feeding areas by concentrating nitrogen near the surface through the release of
flocculent Flocculation, in the field of chemistry, is a process by which colloidal particles come out of suspension to sediment under the form of floc or flake, either spontaneously or due to the addition of a clarifying agent. The action differs from pre ...
fecal Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
plumes. Modified text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
For example, whales and seals may be responsible for replenishing more nitrogen in the
Gulf of Maine The Gulf of Maine is a large gulf of the Atlantic Ocean on the east coast of North America. It is bounded by Cape Cod at the eastern tip of Massachusetts in the southwest and by Cape Sable Island at the southern tip of Nova Scotia in the northeast ...
's euphotic zone than the input of all rivers combined. This upward whale pump played a much larger role before industrial fishing devastated marine mammal stocks, when recycling of nitrogen was likely more than three times the atmospheric nitrogen input. The biological pump mediates the removal of carbon and nitrogen from the euphotic zone through the downward flux of aggregates, feces, and vertical migration of invertebrates and fish. Copepods and other zooplankton produce sinking fecal pellets and contribute to downward transport of dissolved and particulate organic matter by respiring and excreting at depth during migration cycles, thus playing an important role in the export of nutrients (N, P, and Fe) from surface waters. Zooplankton feed in the euphotic zone and export nutrients via sinking fecal pellets, and vertical migration. Fish typically release nutrients at the same depth at which they feed. Excretion for marine mammals, tethered to the surface for respiration, is expected to be shallower in the water column than where they feed. Marine mammals provide important ecosystem services. On a global scale, they can influence climate, through fertilization events and the export of carbon from surface waters to the deep sea through sinking whale carcasses. In coastal areas, whales retain nutrients locally, increasing ecosystem productivity and perhaps raising the carrying capacity for other marine consumers, including commercial fish species. It has been estimated that, in terms of carbon sequestration, one whale is equivalent to thousands of trees.


Vertical migrations

Diel vertically migrating krill, salps, smaller zooplankton and fish can actively transport carbon to depth by consuming POC in the surface layer at night, and metabolising it at their daytime, mesopelagic residence depths. Depending on species life history, active transport may occur on a seasonal basis as well. Without vertical migration the biological pump wouldn't be nearly as efficient. Organisms migrate up to feed at night so when they migrate back to depth during the day they defecate large sinking fecal pellets. Whilst some larger fecal pellets can sink quite fast, the speed that organisms move back to depth is still faster. At night organisms are in the top 100 metres of the water column, but during the day they move down to between 800 and 1000 metres. If organisms were to defecate at the surface it would take the fecal pellets days to reach the depth that they reach in a matter of hours. Therefore, by releasing fecal pellets at depth they have almost 1000 metres less to travel to get to the deep ocean. This is something known as
active transport In cellular biology, ''active transport'' is the movement of molecules or ions across a cell membrane from a region of lower concentration to a region of higher concentration—against the concentration gradient. Active transport requires cellul ...
. The organisms are playing a more active role in moving organic matter down to depths. Because a large majority of the deep sea, especially marine microbes, depends on nutrients falling down, the quicker they can reach the ocean floor the better.
Zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
and
salps A salp (plural salps, also known colloquially as “sea grape”) or salpa (plural salpae or salpas) is a barrel-shaped, planktic tunicate. It moves by contracting, thereby pumping water through its gelatinous body, one of the most efficient ...
play a large role in the active transport of fecal pellets. 15–50% of zooplankton biomass is estimated to migrate, accounting for the transport of 5–45% of particulate organic nitrogen to depth. Salps are large gelatinous plankton that can vertically migrate 800 metres and eat large amounts of food at the surface. They have a very long gut retention time, so fecal pellets usually are released at maximum depth. Salps are also known for having some of the largest fecal pellets. Because of this they have a very fast sinking rate, small
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
particles are known to aggregate on them. This makes them sink that much faster. So while currently there is still much research being done on why organisms vertically migrate, it is clear that vertical migration plays a large role in the active transport of dissolved organic matter to depth.


Lipid pump

The lipid pump sequesters
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent In chemistry, the valence (US spelling) or valency (British spelling) of an element is the measure of its combining capacity with o ...
from the ocean's surface to deeper waters via
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
s associated with
overwintering Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal activi ...
vertically migratory
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
. Lipids are a class of
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and hydrophobic, and their odors are usually weak or ex ...
rich,
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
and
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
deficient compounds essential for cellular structures. The lipid associated carbon enters the
deep ocean The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of 200 metres (656 feet) or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combin ...
as carbon dioxide produced by
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellula ...
of lipid reserves and as organic matter from the mortality of zooplankton. Compared to the more general biological pump, the lipid pump also results in a lipid shunt, where other
nutrient A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excret ...
s like nitrogen and phosphorus that are consumed in excess must be
excreted Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after lea ...
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.
Carbon sequestration Carbon sequestration is the process of storing carbon in a carbon pool. Carbon dioxide () is naturally captured from the atmosphere through biological, chemical, and physical processes. These changes can be accelerated through changes in land ...
via the lipid pump is therefore decoupled from nutrient removal, allowing carbon uptake by oceanic primary production 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. 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
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s of the genus ''
Calanus ''Calanus'' is a genus of marine copepod in the family Calanidae (Order Calanoida). The genus was split in 1974, with some species being placed in a new genus, '' Neocalanus''. The following species are recognised: *'' Calanus aculeatus'' Bra ...
'' in the
Arctic Ocean The Arctic Ocean is the smallest and shallowest of the world's five major oceans. It spans an area of approximately and is known as the coldest of all the oceans. The International Hydrographic Organization (IHO) recognizes it as an ocean, a ...
almost equals that transported below the same depth annually by
particulate organic carbon Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
(POC) in this region. A significant fraction of this transported carbon would not return to the surface due to respiration and mortality. Research is ongoing to more precisely estimate the amount that remains at depth. The export rate of the lipid pump may vary from 1–9.3 g C m−2 y−1 across temperate and subpolar regions containing seasonally-migrating zooplankton. The role of zooplankton, and particularly copepods, in the
food web A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one ...
is crucial to the survival of higher
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...
organisms whose primary source of
nutrition Nutrition is the biochemical and physiological process by which an organism uses food to support its life. It provides organisms with nutrients, which can be metabolized to create energy and chemical structures. Failure to obtain sufficient n ...
is copepods. With warming oceans and increasing melting of
ice cap In glaciology, an ice cap is a mass of ice that covers less than of land area (usually covering a highland area). Larger ice masses covering more than are termed ice sheets. Description Ice caps are not constrained by topographical features ...
s due to
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...
, the organisms associated with the lipid pump may be affected, thus influencing the survival of many commercially important
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
and
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their reli ...
s. As a new and previously unquantified component of oceanic carbon sequestration, further research on the lipid pump can improve the accuracy and overall understanding of carbon fluxes in global oceanic systems.


Bioluminescent shunt

Luminous bacteria Bioluminescent bacteria are light-producing bacteria that are predominantly present in sea water, marine sediments, the surface of decomposing fish and in the gut of marine animals. While not as common, bacterial bioluminescence is also found in t ...
in light organ symbioses are successively acquired by host (squid, fish) from the seawater while they are juveniles, then regularly released into the ocean. Modified text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
In the diagram on the right, depending on the light organ position, luminous bacteria are released from their guts into
fecal pellet Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
s or directly into the seawater (step 1). Motile luminous bacteria colonize organic matter sinking along the
water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...
. Bioluminescent bacteria colonising fecal pellets and particles influence zooplankton consumption rates. Such visual markers increase detection (“bait hypothesis”), attraction and finally predation by upper
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...
s (step 2). In the
mesopelagic The mesopelagic zone (Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins ...
, zooplankton and their predators feed on sinking luminous particles and fecal pellets, which form either aggregates (repackaging) of faster sinking rates or fragment organic matter (due to sloppy feeding) with slower sinking rates (step 3).
Filter feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s also aggregate sinking organic matter without particular visual detection and selection of luminous matter. Diel (and seasonal) vertical migrators feeding on luminous food metabolize and release glowing fecal pellets from the surface to the mesopelagic zone (step 4). This implies bioluminescent bacteria dispersion at large spatial scales, for zooplankton or even some fish actively swimming long distances. Luminous bacteria attached to particles sink down to the seafloor, and sediment can be resuspended by oceanographic physical conditions (step 5) and consumed by epi-benthic organisms. Instruments are (a) plankton net, (b) fish net, (c) Niskin water sampler, (d) bathyphotometer, (e)
sediment trap Sediment traps are instruments used in oceanography and limnology to measure the quantity of sinking particulate organic (and inorganic) material in aquatic systems, usually oceans, lakes, or reservoirs. This flux of material is the product ...
s, (f)
autonomous underwater vehicle An autonomous underwater vehicle (AUV) is a robot that travels underwater without requiring input from an operator. AUVs constitute part of a larger group of undersea systems known as unmanned underwater vehicles, a classification that includes ...
s, (g)
photomultiplier A photomultiplier is a device that converts incident photons into an electrical signal. Kinds of photomultiplier include: * Photomultiplier tube, a vacuum tube converting incident photons into an electric signal. Photomultiplier tubes (PMTs for sho ...
module, (h) astrophysics optical modules
ANTARES Antares is the brightest star in the constellation of Scorpius. It has the Bayer designation α Scorpii, which is Latinised to Alpha Scorpii. Often referred to as "the heart of the scorpion", Antares is flanked by σ Scorpii and τ S ...
and (i–j)
remotely operated vehicle A remotely operated underwater vehicle (technically ROUV or just ROV) is a tethered underwater mobile device, commonly called ''underwater robot''. Definition This meaning is different from remote control vehicles operating on land or in the ai ...
s.


Quantification

The geologic component of the carbon cycle operates slowly in comparison to the other parts of the global carbon cycle. It is one of the most important determinants of the amount of carbon in the atmosphere, and thus of global temperatures. As the biological pump plays an important role in the Earth's carbon cycle, significant effort is spent quantifying its strength. However, because they occur as a result of poorly constrained ecological interactions usually at depth, the processes that form the biological pump are difficult to measure. A common method is to estimate primary production fuelled by
nitrate Nitrate is a polyatomic ion A polyatomic ion, also known as a molecular ion, is a covalent bonded set of two or more atoms, or of a metal complex, that can be considered to behave as a single unit and that has a net charge that is not zer ...
and
ammonium The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaternary a ...
as these nutrients have different sources that are related to the remineralisation of sinking material. From these it is possible to derive the so-called
f-ratio F-ratio or f-ratio may refer to: * The F-ratio used in statistics, which relates the variances of independent samples; see F-distribution * f-ratio (oceanography), which relates recycled and total primary production in the surface ocean * f-number ...
, a proxy for the local strength of the biological pump. Applying the results of local studies to the global scale is complicated by the role the ocean's circulation plays in different ocean regions.


Effects of climate change

Changes in land use, the
combustion Combustion, or burning, is a high-temperature exothermic redox chemical reaction between a fuel (the reductant) and an oxidant, usually atmospheric oxygen, that produces oxidized, often gaseous products, in a mixture termed as smoke. Combusti ...
of
fossil fuel A fossil fuel is a hydrocarbon-containing material formed naturally in the Earth's crust from the remains of dead plants and animals that is extracted and burned as a fuel. The main fossil fuels are coal, oil, and natural gas. Fossil fuels m ...
s, and the production of
cement A cement is a binder, a chemical substance used for construction that sets, hardens, and adheres to other materials to bind them together. Cement is seldom used on its own, but rather to bind sand and gravel ( aggregate) together. Cement mix ...
have led to an increase in CO2 concentration in the atmosphere. At present, about one third (approximately 2 Pg C y−1 = 2 × 1015 grams of carbon per year)Orr, J. C., E. Maier-Reimer, U. Mikolajewicz, P. Monfray, J. L. Sarmiento, J. R. Toggweiler, N. K. Taylor, J. Palmer, N. Gruber, C. L. Sabine, C. Le Quéré, R. M. Key and J. Boutin (2001). Estimates of anthropogenic carbon uptake from four three-dimensional global ocean models. ''Global Biogeochem. Cycles'' 15, 43–60. of anthropogenic emissions of CO2 may be entering the ocean, but this is quite uncertain. Some research suggests that a link between elevated CO2 and marine primary production exists.
Climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...
may affect the biological pump in the future by warming and stratifying the surface ocean. It is believed that this could decrease the supply of nutrients to the euphotic zone, reducing primary production there. Also, changes in the ecological success of calcifying organisms caused by
ocean acidification Ocean acidification is the reduction in the pH value of the Earth’s ocean. Between 1751 and 2021, the average pH value of the ocean surface has decreased from approximately 8.25 to 8.14. The root cause of ocean acidification is carbon dioxid ...
may affect the biological pump by altering the strength of the hard tissues pump. This may then have a "knock-on" effect on the soft tissues pump because calcium carbonate acts to ballast sinking organic material. The second diagram on the right shows some possible effects of sea ice decline and permafrost thaw on Arctic carbon fluxes. On land, plants take up carbon while microorganisms in the soil produce methane and respire CO2. Lakes are net emitters of methane, and organic and inorganic carbon (dissolved and particulate) flow into the ocean through freshwater systems. In the ocean, methane can be released from thawing subsea permafrost, and CO2 is absorbed due to an undersaturation of CO2 in the water compared with the atmosphere. In addition, multiple fluxes are closely associated to sea ice. Current best estimates of atmospheric fluxes are given in Tg C year−1, where available. Note that the emission estimate for lakes is for the area North of ~50º N rather than the narrower definition of arctic tundra for the other terrestrial fluxes. When available, uncertainty ranges are shown in brackets. The arrows do not represent the size of each flux. The biological pump is thought to have played significant roles in atmospheric CO2 fluctuations during past glacial-interglacial periods. However, it is not yet clear how the biological pump will respond to future climate change. For such predictions to be reasonable, it is important to first decipher the response of phytoplankton, one of the key components of the biological pump to future changes in atmospheric CO2. Due to their phylogenetic diversity, different phytoplankton taxa will likely respond to climate change in different ways. For instance, a decrease in the abundance of diatom is expected due to increased stratification in the future ocean. Diatoms are highly efficient in transporting carbon to depths by forming large, rapidly sinking aggregates and their reduced numbers could in turn lead to decreased carbon export. Further, decreased ocean pH due to ocean acidification may thwart the ability of coccolithophores to generate calcareous plates, potentially affecting the biological pump; however, it appears that some species are more sensitive than others. Thus, future changes in the relative abundance of these or other phytoplankton taxa could have a marked impact on total ocean productivity, subsequently affecting ocean biogeochemistry and carbon storage. A 2015 study determined that
coccolithophore Coccolithophores, or coccolithophorids, are single celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdo ...
concentrations in the North Atlantic have increased by an order of magnitude since the 1960s and an increase in absorbed CO2, as well as temperature, were modeled to be the most likely cause of this increase. In a 2017 study, scientists used
species distribution modelling Species distribution modelling (SDM), also known as environmental (or ecological) niche modelling (ENM), habitat modelling, predictive habitat distribution modelling, and range mapping uses computer algorithms to predict the distribution of a spec ...
(SDM) to predict the future global distribution of two phytoplankton species important to the biological pump: the diatom ''
Chaetoceros diadema ''Chaetoceros diadema'' is a diatom in the genus'' Chaetoceros''. The easiest way to identify this species is by finding the very characteristic diadem-like resting spores. Species description Cells united into chains. Very characteristic, diade ...
'' and the coccolithophore ''
Emiliania huxleyi ''Emiliania huxleyi'' is a species of coccolithophore found in almost all ocean ecosystems from the equator to sub-polar regions, and from nutrient rich upwelling zones to nutrient poor oligotrophic waters. It is one of thousands of different ...
''. They employed environmental data described in the
IPCC The Intergovernmental Panel on Climate Change (IPCC) is an intergovernmental body of the United Nations. Its job is to advance scientific knowledge about climate change caused by human activities. The World Meteorological Organization (WMO) a ...
Representative Concentration Pathways scenario 8.5, which predicts
radiative forcing Radiative forcing (or climate forcing) is the change in energy flux in the atmosphere caused by natural or anthropogenic factors of climate change as measured by watts / metre2. It is a scientific concept used to quantify and compare the external ...
in the year 2100 relative to pre-industrial values. Their modelling results predicted that the total ocean area covered by ''C. diadema'' and ''E. huxleyi'' would decline by 8% and 16%, respectively, under the examined climate scenario. They predicted changes in the range and distribution of these two phytoplankton species under these future ocean conditions, if realized, might result in reduced contribution to carbon sequestration via the biological pump. In 2019, a study indicated that at current rates of seawater acidification, we could see Antarctic phytoplanktons smaller and less effective at storing carbon before the end of the century.


Monitoring

{{see also, Remote sensing (oceanography) Monitoring the biological pump is critical to understanding how the Earth's carbon cycle is changing. A variety of techniques are used to monitor the biological pump, which can be deployed from various platforms such as ships, autonomous vehicles, and satellites. At present, satellite remote sensing is the only tool available for viewing the entire surface ocean at high temporal and spatial scales. {{clear


Needed research

Multidisciplinary observations are still needed in the deep
water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...
to properly understand the biological pump:Levin, L.A., Bett, B.J., Gates, A.R., Heimbach, P., Howe, B.M., Janssen, F., McCurdy, A., Ruhl, H.A., Snelgrove, P., Stocks, K.I., Bailey, D. and 27 others (2019) "Global observing needs in the deep ocean". ''Frontiers in Marine Science'', 6: 241. {{doi, 10.3389/fmars.2019.00241. Modified text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
* Physics: stratification affects particle sinking; understanding the origin of the particles and the residence time of the DIC from particle remineralization in the deep ocean requires measurement of advection and mixing. * Biogeochemistry: export/mixing down of particulate and dissolved organic matter from the surface layer determines labile organic matter arriving at the seafloor, which is either respired by seafloor biota or stored for longer times in the sediment. * Biology and ecosystems: zooplankton and microorganisms break down and remineralize sinking particles in the water column. Exported organic matter feeds all water column and benthic biota (zooplankton, benthic invertebrates, microbes) sustaining their biomass, density, and biodiversity.


See also

*
f-ratio (oceanography) In oceanic biogeochemistry, the f-ratio is the fraction of total primary production fuelled by nitrate (as opposed to that fuelled by other nitrogen compounds such as ammonium). The ratio was originally defined by Richard Eppley and Bruce Pe ...
*
Lysocline The lysocline is the depth in the ocean dependent upon the carbonate compensation depth (CCD), usually around 3.5 km, below which the rate of dissolution of calcite increases dramatically because of a pressure effect. While the lysocline is the up ...
*
Mooring (oceanography) current_meter.html" ;"title="sediment traps and current meter">sediment traps and current meters --> A mooring in oceanography is a collection of devices connected to a wire and anchored on the sea floor. It is the ''Lagrangian and Eulerian speci ...
*
Apparent oxygen utilisation In freshwater or marine systems apparent oxygen utilization (AOU) is the difference between oxygen gas solubility (i.e. the concentration at saturation) and the measured oxygen concentration in water with the same physical and chemical properties ...
{{clear


References

{{reflist {{DEFAULTSORT:Biological Pump Aquatic ecology Biological oceanography Carbon cycle Chemical oceanography