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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
micrometer Micrometer can mean: * Micrometer (device), used for accurate measurements by means of a calibrated screw * American spelling of micrometre The micrometre ( international spelling as used by the International Bureau of Weights and Measures; ...
s. The fraction remaining on the filter is called particulate organic carbon (POC). Dissolved organic matter (DOM) is a closely related term often used interchangeably with DOC. While DOC refers specifically to the mass of carbon in the dissolved organic material, DOM refers to the total mass of the dissolved organic matter. So DOM also includes the mass of other elements present in the organic material, such as nitrogen, oxygen and hydrogen. DOC is a component of DOM and there is typically about twice as much DOM as DOC. Many statements that can be made about DOC apply equally to DOM, and ''vice versa''. DOC is abundant in
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military ...
and
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does incl ...
systems and is one of the greatest cycled reservoirs 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 ...
on Earth, accounting for the same amount of
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent—its atom making four electrons available to form covalent chemical bonds. It belongs to group 14 of the periodic table. Carbon ma ...
as in the atmosphere and up to 20% of all organic carbon. In general, organic carbon compounds are the result of
decomposition Decomposition or rot is the process by which dead organic substances are broken down into simpler organic or inorganic matter such as carbon dioxide, water, simple sugars and mineral salts. The process is a part of the nutrient cycle and ...
processes from dead organic matter including plants and animals. DOC can originate from within or outside any given body of water. DOC originating from within the body of water is known as autochthonous DOC and typically comes from
aquatic plant Aquatic plants are plants that have adapted to living in aquatic environments ( saltwater or freshwater). They are also referred to as hydrophytes or macrophytes to distinguish them from algae and other microphytes. A macrophyte is a plant that ...
s or
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
, while DOC originating outside the body of water is known as allochthonous DOC and typically comes from
soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former ...
s or terrestrial plants. When water originates from land areas with a high proportion of organic soils, these components can drain into rivers and lakes as DOC. The marine DOC pool is important for the functioning of marine ecosystems because they are at the interface between the chemical and the biological worlds. DOC fuels
marine food webs Compared to terrestrial environments, marine environments have biomass pyramids which are inverted at the base. In particular, the biomass of consumers (copepods, krill, shrimp, forage fish) is larger than the biomass of primary producers. Thi ...
, and is a major component of the Earth's carbon cycling.Lønborg, C., Carreira, C., Jickells, T. and Álvarez-Salgado, X.A. (2020) "Impacts of global change on ocean dissolved organic carbon (DOC) cycling". ''Frontiers in Marine Science'', 7: 466. . Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License


Overview

DOC is a basic nutrient, supporting growth of
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s and plays an important role in the global
carbon cycle 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 compon ...
through the microbial loop. In some organisms (stages) that do not feed in the traditional sense, dissolved matter may be the only external food source. Moreover, DOC is an indicator of organic loadings in streams, as well as supporting terrestrial processing (e.g., within soil, forests, and wetlands) of organic matter. Dissolved organic carbon has a high proportion of biodegradable dissolved organic carbon (BDOC) in first order streams compared to higher order streams. In the absence of extensive
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (Anoxic waters, anoxic) processes prevailing, especially in t ...
s,
bog A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg; a ...
s, or
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
s, baseflow concentrations of DOC in undisturbed watersheds generally range from approximately 1 to 20 mg/L carbon. Carbon concentrations considerably vary across ecosystems. For example, the
Everglades The Everglades is a natural region of tropical wetlands in the southern portion of the U.S. state of Florida, comprising the southern half of a large drainage basin within the Neotropical realm. The system begins near Orlando with the Kissim ...
may be near the top of the range and the middle of oceans may be near the bottom. Occasionally, high concentrations of organic carbon indicate anthropogenic influences, but most DOC originates naturally. The ''BDOC fraction'' consists of organic
molecule 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 b ...
s that
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 ...
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 am ...
can use as a source of energy and carbon. Some subset of DOC constitutes the precursors of disinfection byproducts for drinking water. BDOC can contribute to undesirable biological regrowth within water distribution systems. The dissolved fraction of
total 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 organi ...
(TOC) is an operational classification. Many researchers use the term "dissolved" for compounds that pass through a 0.45 μm filter, but 0.22 μm filters have also been used to remove higher colloidal concentrations. A practical definition of dissolved typically used in
marine chemistry Marine chemistry, also known as ocean chemistry or chemical oceanography, is influenced by plate tectonics and seafloor spreading, turbidity currents, sediments, pH levels, atmospheric constituents, metamorphic activity, and ecology. The fiel ...
is all substances that pass through a GF/F filter, which has a nominal pore size of approximately 0.7 μm (Whatman glass microfiber filter, 0.6–0.8 μm particle retention). The recommended procedure is the HTCO technique, which calls for filtration through pre-combusted glass fiber filters, typically the GF/F classification.


Labile and recalcitrant

Dissolved organic matter can be classified as labile or as recalcitrant, depending on its reactivity. Recalcitrant DOC is also called
refractory In materials science, a refractory material or refractory is a material that is resistant to decomposition by heat, pressure, or chemical attack, and retains strength and form at high temperatures. Refractories are polycrystalline, polyphase, ...
DOC, and these terms seem to be used interchangeably in the context of DOC. Depending on the origin and composition of DOC, its behavior and cycling are different; the labile fraction of DOC decomposes rapidly through microbially or photochemically mediated processes, whereas refractory DOC is resistant to degradation and can persist in the ocean for millennia. In the coastal ocean, organic matter from terrestrial plant litter or soils appears to be more refractory and thus often behaves conservatively. In addition, refractory DOC is produced in the ocean by the bacterial transformation of labile DOC, which reshapes its composition. Due to the continuous production and degradation in natural systems, the DOC pool contains a spectrum of reactive compounds each with their own reactivity,Vahatalo, A. V., Aarnos, H., and Mantyniemi, S. (2010). Biodegradability continuum and biodegradation kinetics of natural organic matter described by the beta distribution. Biogeochemistry 100, 227–240. doi: 10.1007/s10533-010-9419-4 that have been divided into fractions from labile to recalcitrant, depending on the turnover times, as shown in the following table... This wide range in turnover or degradation times has been linked with the chemical composition, structure and molecular size,Amon, R. M. W., and Benner, R. (1996). Bacterial utilization of different size classes of dissolved organic matter. Limnol. Oceanogr. 41, 41–51. doi: 10.4319/lo.1996.41.1.0041 but degradation also depends on the environmental conditions (e.g., nutrients), prokaryote diversity, redox state, iron availability, mineral-particle associations, temperature, sun-light exposure, biological production of recalcitrant compounds, and the effect of priming or dilution of individual molecules. For example, lignin can be degraded in aerobic soils but is relatively recalcitrant in anoxic marine sediments. This example shows bioavailability varies as a function of the ecosystem's properties. Accordingly, even normally ancient and recalcitrant compounds, such as petroleum, carboxyl-rich alicyclic molecules, can be degraded in the appropriate environmental setting.


Terrestrial ecosystems


Soil

Dissolved organic matter (DOM) is one of the most active and mobile carbon pools and has an important role in global carbon cycling. In addition, dissolved organic carbon (DOC) affects the soil negative electrical charges
denitrification Denitrification is a microbially facilitated process where nitrate (NO3−) is reduced and ultimately produces molecular nitrogen (N2) through a series of intermediate gaseous nitrogen oxide products. Facultative anaerobic bacteria perform denit ...
process, acid-base reactions in the soil solution, retention and translocation of nutrients (
cation An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by conven ...
s), and immobilization of
heavy metals upright=1.2, Crystals of osmium, a heavy metal nearly twice as dense as lead">lead.html" ;"title="osmium, a heavy metal nearly twice as dense as lead">osmium, a heavy metal nearly twice as dense as lead Heavy metals are generally defined as ...
and
xenobiotic A xenobiotic is a chemical substance found within an organism that is not naturally produced or expected to be present within the organism. It can also cover substances that are present in much higher concentrations than are usual. Natural compo ...
s. Soil DOM can be derived from different sources (inputs), such as atmospheric carbon dissolved in rainfall, litter and crop residues, manure, root exudates, and decomposition of soil organic matter (SOM). In the soil, DOM availability depends on its interactions with mineral components (e.g., clays, Fe and Al oxides) modulated by
adsorption Adsorption is the adhesion of atoms, ions or molecules from a gas, liquid or dissolved solid to a surface. This process creates a film of the ''adsorbate'' on the surface of the ''adsorbent''. This process differs from absorption, in which a ...
and
desorption Desorption is the physical process where a previously adsorbed substance is released from a surface. This happens when a molecule gains enough energy to overcome the activation barrier of the bounding energy that keeps it in the surface. There ...
processes.Saidy, A.R.; Smernik, R.J.; Baldock, J.A.; Kaiser, K.; Sanderman, J. 2015. Microbial degradation of organic carbon sorbed to phyllosilicate clays with and without hydrous iron oxide coating. European Journal of Soil Science 66: 83–94. It also depends on SOM fractions (e.g., stabilized organic molecules and microbial biomass) by mineralization and immobilization processes. In addition, the intensity of these interactions changes according to soil inherent properties, land use, and crop management. During the decomposition of organic material, most carbon is lost as CO2 to the atmosphere by microbial oxidation.
Soil type A soil type is a taxonomic unit in soil science. All soils that share a certain set of well-defined properties form a distinctive soil type. Soil type is a technical term of soil classification, the science that deals with the systematic categori ...
and landscape slope, leaching, and
runoff Runoff, run-off or RUNOFF may refer to: * RUNOFF, the first computer text-formatting program * Runoff or run-off, another name for bleed, printing that lies beyond the edges to which a printed sheet is trimmed * Runoff or run-off, a stock marke ...
are also important processes associated to DOM losses in the soil.Veum, K.S.; Goyne, K.W.; Motavalli, P.P.; Udawatta, R.P. 2009. Runoff and dissolved organic carbon loss from a paired-watershed study of three adjacent agricultural Watersheds. Agriculture, Ecosystems & Environment 130: 115–122. In well-drained soils, leached DOC can reach the
water table The water table is the upper surface of the zone of saturation. The zone of saturation is where the pores and fractures of the ground are saturated with water. It can also be simply explained as the depth below which the ground is saturated. T ...
and release nutrients and pollutants that can contaminate
groundwater Groundwater is the water present beneath Earth's surface in rock and Pore space in soil, soil pore spaces and in the fractures of stratum, rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit ...
, whereas runoff transports DOM and
xenobiotic A xenobiotic is a chemical substance found within an organism that is not naturally produced or expected to be present within the organism. It can also cover substances that are present in much higher concentrations than are usual. Natural compo ...
s to other areas, rivers, and lakes.


Groundwater

Precipitation and surface water leaches dissolved organic carbon (DOC) from vegetation and
plant litter Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that have fallen to the ground. This detritus or dead organic material and its constitue ...
and percolates through the soil column to the saturated zone. The concentration, composition, and bioavailability of DOC are altered during transport through the soil column by various physicochemical and biological processes, including
sorption Sorption is a physical and chemical process by which one substance becomes attached to another. Specific cases of sorption are treated in the following articles: ; Absorption: "the incorporation of a substance in one state into another of a di ...
,
desorption Desorption is the physical process where a previously adsorbed substance is released from a surface. This happens when a molecule gains enough energy to overcome the activation barrier of the bounding energy that keeps it in the surface. There ...
,
biodegradation Biodegradation is the breakdown of organic matter by microorganisms, such as bacteria and fungi. It is generally assumed to be a natural process, which differentiates it from composting. Composting is a human-driven process in which biodegra ...
and
biosynthesis Biosynthesis is a multi-step, enzyme-catalyzed process where substrates are converted into more complex products in living organisms. In biosynthesis, simple compounds are modified, converted into other compounds, or joined to form macromolecul ...
.
Hydrophobic In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, ...
molecules are preferentially partitioned onto soil minerals and have a longer retention time in soils than
hydrophilic A hydrophile is a molecule or other molecular entity that is attracted to water molecules and tends to be dissolved by water.Liddell, H.G. & Scott, R. (1940). ''A Greek-English Lexicon'' Oxford: Clarendon Press. In contrast, hydrophobes are n ...
molecules. The hydrophobicity and retention time of colloids and dissolved molecules in soils are controlled by their size, polarity, charge, and
bioavailability In pharmacology, bioavailability is a subcategory of absorption and is the fraction (%) of an administered drug that reaches the systemic circulation. By definition, when a medication is administered intravenously, its bioavailability is 100%. Ho ...
. Bioavailable DOM is subjected to microbial decomposition, resulting in a reduction in size and molecular weight. Novel molecules are synthesized by
soil microbe Soil microbiology is the study of microorganisms in soil, their functions, and how they affect soil properties. It is believed that between two and four billion years ago, the first ancient bacteria and microorganisms came about on Earth's oceans. ...
s, and some of these metabolites enter the DOC reservoir in groundwater.Shen, Y., Chapelle, F.H., Strom, E.W. and Benner, R. (2015) "Origins and bioavailability of dissolved organic matter in groundwater". ''Biogeochemistry'', 122(1): 61–78. . Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License


Freshwater ecosystems

Aquatic carbon occurs in different forms. Firstly, a division is made between organic and inorganic carbon. Organic carbon is a mixture of organic compounds originating from detritus or primary producers. It can be divided into POC ( particulate organic carbon; particles > 0.45 μm) and DOC (dissolved organic carbon; particles < 0.45 μm). DOC usually makes up 90% of the total amount of aquatic organic carbon. Its concentration ranges from 0.1 to >300 mg L−1. Likewise, inorganic carbon also consists of a particulate (PIC) and a dissolved phase (DIC). PIC mainly consists of
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 ...
s (e.g., CaCO3), DIC consists of carbonate (CO32-),
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 biochemi ...
(HCO3), CO2 and a negligibly small fraction of carbonic acid (H2CO3). The inorganic carbon compounds exist in equilibrium that depends on the pH of the water. DIC concentrations in freshwater range from about zero in acidic waters to 60 mg C L−1 in areas with carbonate-rich sediments. POC can be degraded to form DOC; DOC can become POC by
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 ...
. Inorganic and organic carbon are linked through
aquatic organism An aquatic animal is any animal, whether invertebrate or vertebrate, that lives in water for most or all of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic anim ...
s. CO2 is used in
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 in ...
(P) by for instance macrophytes, 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 cellul ...
(R), and exchanged with the atmosphere. Organic carbon is produced by organisms and is released during and after their life; e.g., in rivers, 1–20% of the total amount of DOC is produced by macrophytes. Carbon can enter the system from the catchment and is transported to the oceans by rivers and streams. There is also exchange with carbon in the sediments, e.g., burial of organic carbon, which is important for
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 lan ...
in aquatic habitats. Aquatic systems are very important in global carbon sequestration; e.g., when different European ecosystems are compared, inland aquatic systems form the second largest carbon sink (19–41 Tg C y−1); only forests take up more carbon (125–223 Tg C y−1).


Marine ecosystems


Sources

In marine systems DOC originates from either autochthonous or allochthonous sources. Autochthonous DOC is produced within the system, primarily by plankton organisms Kawasaki, N., and Benner, R. (2006). Bacterial release of dissolved organic matter during cell growth and decline: molecular origin and composition. Limnol. Oceanogr. 51, 2170–2180. doi: 10.4319/lo.2006.51.5.2170 and in coastal waters additionally by benthic microalgae, benthic fluxes, and macrophytes, whereas allochthonous DOC is mainly of terrestrial origin supplemented by groundwater and atmospheric inputs.Raymond, P. A., and Spencer, R. G. M. (2015). "Riverine DOM", in Biogeochemistry of Marine Dissolved Organic Matter, eds D. A. Hansell and C. A. Carlson (Amsterdam: Elsevier), 509–533. doi: 10.1016/B978-0-12-405940-5.00011-X In addition to soil derived humic substances, terrestrial DOC also includes material leached from plants exported during rain events, emissions of plant materials to the atmosphere and deposition in aquatic environments (e.g., volatile organic carbon and pollens), and also thousands of synthetic human-made organic chemicals that can be measured in the ocean at trace concentrations. Dissolved organic carbon (DOC) represents one of the Earth's major carbon pools. It contains a similar amount of carbon as the atmosphere and exceeds the amount of carbon bound in marine biomass by more than two-hundred times. DOC is mainly produced in the near-surface layers 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 ...
and zooplankton grazing processes. Other sources of marine DOC are dissolution from particles, terrestrial and
hydrothermal vent A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspo ...
input, and microbial production.
Prokaryotes A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Con ...
(bacteria and archaea) contribute to the DOC pool via release of capsular material, exopolymers, and hydrolytic enzymes, as well as via mortality (e.g.
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 recy ...
). Prokaryotes are also the main decomposers of DOC, although for some of the most recalcitrant forms of DOC very slow abiotic degradation in hydrothermal systems or possibly sorption to sinking particles may be the main removal mechanism. Mechanistic knowledge about DOC-microbe-interactions is crucial to understand the cycling and distribution of this active carbon reservoir. Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License


Phytoplankton

Phytoplankton produces DOC by
extracellular This glossary of biology terms is a list of definitions of fundamental terms and concepts used in biology, the study of life and of living organisms. It is intended as introductory material for novices; for more specific and technical definitions ...
release commonly accounting between 5 and 30% of their total primary production, although this varies from species to species. Nonetheless, this release of extracellular DOC is enhanced under high light and low nutrient levels, and thus should increase relatively from eutrophic to oligotrophic areas, probably as a mechanism for dissipating cellular energy. Phytoplankton can also produce DOC by autolysis during physiological stress situations e.g., nutrient limitation. Other studies have demonstrated DOC production in association with meso- and macro-zooplankton feeding on phytoplankton and bacteria.Hygum, B. H., Petersen, J. W., and Søndergaard, M. (1997). Dissolved organic carbon released by zooplankton grazing activity- a high quality substrate pool for bacteria. J. Plankton Res. 19, 97–111. doi: 10.1093/plankt/19.1.97


Zooplankton

Zooplankton-mediated release of DOC occurs through sloppy feeding, excretion and defecation which can be important energy sources for microbes. Such DOC production is largest during periods with high food concentration and dominance of large zooplankton species.


Bacteria and viruses

Bacteria are often viewed as the main consumers of DOC, but they can also produce DOC during
cell division Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there ...
and
viral 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 ...
. The biochemical components of bacteria are largely the same as other organisms, but some compounds from the cell wall are unique and are used to trace bacterial derived DOC (e.g.,
peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ba ...
). These compounds are widely distributed in the ocean, suggesting that bacterial DOC production could be important in marine systems. Viruses are the most abundant life forms in the oceans infecting all life forms including algae, bacteria and zooplankton. After infection, the virus either enters a dormant (
lysogenic Lysogeny, or the lysogenic cycle, is one of two cycles of viral reproduction (the lytic cycle being the other). Lysogeny is characterized by integration of the bacteriophage nucleic acid into the host bacterium's genome or formation of a circu ...
) or productive (
lytic The lytic cycle ( ) is one of the two cycles of viral reproduction (referring to bacterial viruses or bacteriophages), the other being the lysogenic cycle. The lytic cycle results in the destruction of the infected cell and its membrane. Bacter ...
) state. The lytic cycle causes disruption of the cell(s) and release of DOC.


Macrophytes

Marine macrophytes (i.e.,
macroalgae Seaweed, or macroalgae, refers to thousands of species of macroscopic, multicellular, marine algae. The term includes some types of '' Rhodophyta'' (red), ''Phaeophyta'' (brown) and ''Chlorophyta'' (green) macroalgae. Seaweed species such as ...
and
seagrass Seagrasses are the only flowering plants which grow in marine environments. There are about 60 species of fully marine seagrasses which belong to four families ( Posidoniaceae, Zosteraceae, Hydrocharitaceae and Cymodoceaceae), all in the ...
) are highly productive and extend over large areas in coastal waters but their production of DOC has not received much attention. Macrophytes release DOC during growth with a conservative estimate (excluding release from decaying tissues) suggesting that macroalgae release between 1-39% of their gross primary production, while seagrasses release less than 5% as DOC of their gross primary production. The released DOC has been shown to be rich in carbohydrates, with rates depending on temperature and light availability.Barrón, C., and Duarte, C. M. (2015). Dissolved organic carbon pools and export from the coastal ocean. Glob. Biogeochem. Cycles 29, 1725–1738. doi: 10.1002/2014GB005056 Globally the macrophyte communities have been suggested to produce ~160 Tg C yr−1 of DOC, which is approximately half the annual global river DOC input (250 Tg C yr−1).


Marine sediments

Marine sediment 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, mai ...
s represent the main sites of OM degradation and burial in the ocean, hosting microbes in densities up to 1000 times higher than found in 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 ...
. The DOC concentrations in sediments are often an order of magnitude higher than in the overlying water column. This concentration difference results in a continued diffusive flux and suggests that sediments are a major DOC source releasing 350 Tg C yr−1, which is comparable to the input of DOC from rivers. This estimate is based on calculated diffusive fluxes and does not include resuspension events which also releases DOC and therefore the estimate could be conservative. Also, some studies have shown that geothermal systems and petroleum seepage contribute with pre-aged DOC to the deep
ocean basin In hydrology, an oceanic basin (or ocean basin) is anywhere on Earth that is covered by seawater. Geologically, ocean basins are large  geologic basins that are below sea level. Most commonly the ocean is divided into basins fol ...
s, but consistent global estimates of the overall input are currently lacking. Globally,
groundwater Groundwater is the water present beneath Earth's surface in rock and Pore space in soil, soil pore spaces and in the fractures of stratum, rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit ...
s account for an unknown part of the freshwater DOC flux to the oceans. The DOC in groundwater is a mixture of terrestrial, infiltrated marine, and in situ microbially produced material. This flux of DOC to coastal waters could be important, as concentrations in groundwater are generally higher than in coastal seawater, but reliable global estimates are also currently lacking.


Sinks

The main processes that remove DOC from the ocean water column are: (1) Thermal degradation in e.g., submarine hydrothermal systems;Lang, S. Q., Butterfield, D. A., Lilley, M. D., Paul Johnson, H., and Hedges, J. I. (2006). Dissolved organic carbon in ridge-axis and ridge-flank hydrothermal systems. Geochim. Cosmochim. Acta 70, 3830–3842. doi: 10.1016/j.gca.2006.04.031 (2) bubble
coagulation Coagulation, also known as clotting, is the process by which blood changes from a liquid to a gel, forming a blood clot. It potentially results in hemostasis, the cessation of blood loss from a damaged vessel, followed by repair. The mechanism ...
and
abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...
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 ...
into
microparticle Microparticles are particles between 0.1 and 100 μm in size. Commercially available microparticles are available in a wide variety of materials, including ceramics, glass, polymers, and metals. Microparticles encountered in daily life incl ...
s or
sorption Sorption is a physical and chemical process by which one substance becomes attached to another. Specific cases of sorption are treated in the following articles: ; Absorption: "the incorporation of a substance in one state into another of a di ...
to particles; (3) abiotic degradation via
photochemical reaction Organic photochemistry encompasses organic reactions that are induced by the action of light. The absorption of ultraviolet light by organic molecules often leads to reactions. In the earliest days, sunlight was employed, while in more modern times ...
s;Moran, M. A., and Zepp, R. G. (1997). Role of photoreactions in the formation of biologically labile compounds from dissolved organic matter. Limnol. Oceanogr. 42, 1307–1316. doi: 10.4319/lo.1997.42.6.1307Mopper, K., Kieber, D. J., and Stubbins, A. (2015). "Marine photochemistry of organic matter", in Biogeochemistry of Marine Dissolved Organic Matter, eds C. A. Carlson and D. A. Hansell (Amsterdam: Elsevier), 389–450. doi: 10.1016/B978-0-12-405940-5.00008-X and (4) biotic degradation 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 ...
marine prokaryotes 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 cellula ...
. It has been suggested that the combined effects of photochemical and microbial degradation represent the major sinks of DOC.


Thermal degradation

File:Removal of refractory DOC in the ocean.webp, upright=2.4,
Thermal degradation Thermal decomposition, or thermolysis, is a chemical decomposition caused by heat. The decomposition temperature of a substance is the temperature at which the substance chemically decomposes. The reaction is usually endothermic as heat is req ...
of DOC has been found at high-temperature hydrothermal ridge-flanks, where outflow DOC concentrations are lower than in the inflow. While the global impact of these processes has not been investigated, current data suggest it is a minor DOC sink. Abiotic DOC flocculation is often observed during rapid (minutes) shifts in salinity when fresh and marine waters mix.Sholkovitz, E. R. (1976). Flocculation of dissolved organic and inorganic matter during the mixing of river water and seawater. Geochim. Cosmochim. Acta 40, 831–845. doi: 10.1016/0016-7037(76)90035-1 Flocculation changes the DOC chemical composition, by removing
humic Humic substances (HS) are organic compounds that are important components of humus, the major organic fraction of soil, peat, and coal (and also a constituent of many upland streams, dystrophic lakes, and ocean water). For a long era in the 19th an ...
compounds and reducing molecular size, transforming DOC to particulate organic flocs which can sediment and/or be consumed by grazers and
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, but it also stimulates the bacterial degradation of the flocculated DOC. The impacts of flocculation on the removal of DOC from coastal waters are highly variable with some studies suggesting it can remove up to 30% of the DOC pool, while others find much lower values (3–6%;). Such differences could be explained by seasonal and system differences in the DOC chemical composition, pH, metallic cation concentration, microbial reactivity, and ionic strength.


CDOM

The colored fraction of DOC (CDOM) absorbs light in the blue and UV-light range and therefore influences plankton productivity both negatively by absorbing light, that otherwise would be available for photosynthesis, and positively by protecting plankton organisms from harmful UV-light. However, as the impact of UV damage and ability to repair is extremely variable, there is no consensus on how UV-light changes might impact overall plankton communities. The CDOM absorption of light initiates a complex range of photochemical processes, which can impact nutrient, trace metal and DOC chemical composition, and promote DOC degradation.


Photodegradation

Photodegradation Photodegradation is the alteration of materials by light. Commonly, the term is used loosely to refer to the combined action of sunlight and air, which cause oxidation and hydrolysis. Often photodegradation is intentionally avoided, since it destro ...
involves the transformation of CDOM into smaller and less colored molecules (e.g., organic acids), or into inorganic carbon (CO, CO2), and nutrient salts (NH4, HPO). Therefore, it generally means that photodegradation transforms recalcitrant into labile DOC molecules that can be rapidly used by prokaryotes for biomass production and respiration. However, it can also increase CDOM through the transformation of compounds such as triglycerides, into more complex aromatic compounds, which are less degradable by microbes. Moreover, UV radiation can produce e.g., reactive oxygen species, which are harmful to microbes. The impact of photochemical processes on the DOC pool depends also on the chemical composition, with some studies suggesting that recently produced autochthonous DOC becomes less bioavailable while allochthonous DOC becomes more bioavailable to prokaryotes after sunlight exposure, albeit others have found the contrary. Photochemical reactions are particularly important in coastal waters which receive high loads of terrestrial derived CDOM, with an estimated ~20–30% of terrestrial DOC being rapidly photodegraded and consumed. Global estimates also suggests that in marine systems photodegradation of DOC produces ~180 Tg C yr−1 of inorganic carbon, with an additional 100 Tg C yr−1 of DOC made more available to microbial degradation. Another attempt at global ocean estimates also suggest that photodegradation (210 Tg C yr−1) is approximately the same as the annual global input of riverine DOC (250 Tg C yr−1;), while others suggest that direct photodegradation exceeds the riverine DOC inputs.


Recalcitrant DOC

DOC is conceptually divided into labile DOC, which is rapidly taken up by heterotrophic microbes, and the recalcitrant DOC reservoir, which has accumulated in the ocean (following a definition by Hansell). As a consequence of its recalcitrance, the accumulated DOC reaches average radiocarbon ages between 1,000 and 4,000 years in surface waters, and between 3,000 and 6,000 years in the deep ocean, indicating that it persists through several deep ocean mixing cycles between 300 and 1,400 years each. Behind these average radiocarbon ages, a large spectrum of ages is hidden. Follett et al. showed DOC comprises a fraction of modern radiocarbon age, as well as DOC reaching radiocarbon ages of up to 12,000 years.


Distribution

More precise measurement techniques developed in the late 1990s have allowed for a good understanding of how dissolved organic carbon is distributed in marine environments both vertically and across the surface. It is now understood that dissolved organic carbon in the ocean spans a range from very labile to very recalcitrant (refractory). The labile dissolved organic carbon is mainly produced by marine organisms and is consumed in the surface ocean, and consists of sugars, proteins, and other compounds that are easily used by marine bacteria. Recalcitrant dissolved organic carbon is evenly spread throughout the water column and consists of high molecular weight and structurally complex compounds that are difficult for marine organisms to use such as the
lignin Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidity a ...
,
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
, or
humic acids Humic substances (HS) are organic compounds that are important components of humus, the major organic fraction of soil, peat, and coal (and also a constituent of many upland streams, dystrophic lakes, and ocean water). For a long era in the 19th a ...
. As a result, the observed vertical distribution consists of high concentrations of labile DOC in the upper water column and low concentrations at depth. File:Environmental processes controlling the recalcitrance of oceanic DOC.jpg, The dots represent DOC molecules and arrows represent physicochemical and biological processes that impact DOC concentration and molecular composition. In the surface ocean, DOC derived from primary production is rapidly remineralized or transformed through microbial degradation (black arrow), photochemical degradation (yellow arrow), or particle exchange (green arrow). Labile components are removed down the water column and DOC becomes diluted by processes, such as particle exchange (brown arrow), sediment dissolution (gray arrow), and microbial reworking (white arrow), which continue to alter, add, and/or remove molecules from the bulk DOC pool. Thus, the apparent recalcitrance of DOC in the ocean’s interior is an emergent property that is largely controlled by environmental context. In addition to vertical distributions, horizontal distributions have been modeled and sampled as well. In the surface ocean at a depth of 30 meters, the higher dissolved organic carbon concentrations are found in the South Pacific Gyre, the South Atlantic Gyre, and the Indian Ocean. At a depth of 3,000 meters, highest concentrations are in the North Atlantic Deep Water where dissolved organic carbon from the high concentration surface ocean is removed to depth. While in the northern Indian Ocean high DOC is observed due to high fresh water flux and sediments. Since the time scales of horizontal motion along the ocean bottom are in the thousands of years, the refractory dissolved organic carbon is slowly consumed on its way from the North Atlantic and reaches a minimum in the North Pacific.


As emergent

Dissolved organic matter is a heterogeneous pool of thousands, likely millions, of organic compounds. These compounds differ not only in composition and concentration (from pM to μM), but also originate from various organisms (phytoplankton, zooplankton, and bacteria) and environments (terrestrial vegetation and soils, coastal fringe ecosystems) and may have been produced recently or thousands of years ago. Moreover, even organic compounds deriving from the same source and of the same age may have been subjected to different processing histories prior to accumulating within the same pool of DOM. Interior ocean DOM is a highly modified fraction that remains after years of exposure to sunlight, utilization by heterotrophs, flocculation and coagulation, and interaction with particles. Many of these processes within the DOM pool are compound- or class-specific. For example, condensed aromatic compounds are highly photosensitive, whereas proteins, carbohydrates, and their monomers are readily taken up by bacteria. Microbes and other consumers are selective in the type of DOM they utilize and typically prefer certain organic compounds over others. Consequently, DOM becomes less reactive as it is continually reworked. Said another way, the DOM pool becomes less labile and more refractory with degradation. As it is reworked, organic compounds are continually being added to the bulk DOM pool by physical mixing, exchange with particles, and/or production of organic molecules by the consumer community. As such, the compositional changes that occur during degradation are more complex than the simple removal of more labile components and resultant accumulation of remaining, less labile compounds. Dissolved organic matter recalcitrance (i.e., its overall reactivity toward degradation and/or utilization) is therefore an emergent property. The perception of DOM recalcitrance changes during organic matter degradation and in conjunction with any other process that removes or adds organic compounds to the DOM pool under consideration. The surprising resistance of high concentrations of DOC to microbial degradation has been addressed by several hypotheses. The prevalent notion is that the recalcitrant fraction of DOC has certain chemical properties, which prevent decomposition by microbes ("intrinsic stability hypothesis"). An alternative or additional explanation is given by the "dilution hypothesis", that all compounds are labile, but exist in concentrations individually too low to sustain microbial populations but collectively form a large pool. The dilution hypothesis has found support in recent experimental and theoretical studies.


DOM isolation and analysis

DOM is found in low concentrations in nature for direct analysis with NMR or MS. Moreover, DOM samples often contain high concentrations of inorganic salts that are incompatible with such techniques. Therefore, it is necessary a concentration and isolation step of the sample. The most used isolation techniques are
ultrafiltration Ultrafiltration (UF) is a variety of membrane filtration in which forces such as pressure or concentration gradients lead to a separation through a semipermeable membrane. Suspended solids and solutes of high molecular weight are retained in the ...
,
reverse osmosis Reverse osmosis (RO) is a water purification process that uses a partially permeable membrane to separate ions, unwanted molecules and larger particles from drinking water. In reverse osmosis, an applied pressure is used to overcome osmotic ...
, and solid-phase extraction. Among them solid-phase extraction is considered as the cheapest and easiest technique.


See also

*
Blackwater river A blackwater river is a type of river with a slow-moving channel flowing through forested swamps or wetlands. As vegetation decays, tannins leach into the water, making a transparent, acidic water that is darkly stained, resembling black tea ...
* Dissolved inorganic carbon *
Foam line Foam lines are accumulations of foam, with trapped insects, exuvia, etc. found on the surface of freshwater water courses or bodies.
* Microbial loop *
Total 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 organi ...


References


External links

* Hansell DA and Carlson CA (Eds.) (2014
''Biogeochemistry of Marine Dissolved Organic Matter''
Second edition, Academic Press. . * {{DEFAULTSORT:Dissolved Organic Carbon Environmental chemistry Water quality indicators Water chemistry