Chemocline

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A chemocline is a type of cline, a layer of fluid with different properties, characterized by a strong, vertical chemistry gradient within a body of water. In bodies of water where chemoclines occur, the cline separates the upper and lower layers, resulting in different properties for those layers. [1] The lower layer shows a change in the concentration of dissolved gases and solids compared to the upper layer. [2]

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Chemoclines most commonly occur where local conditions favor the formation of anoxic bottom water deep water deficient in oxygen, where only anaerobic forms of life can exist. Common anaerobic organisms that live in these conditions include phototrophic purple sulfur bacteria and green sulfur bacteria. [3] The Black Sea is an example of a body of water with a prominent chemocline, though similar bodies (classified as meromictic lakes) exist across the globe. [4] [5] Meromictic lakes are the result of meromixis, which is a circumstance where a body of water does not fully mix and circulate, causing stratification. [1] [6]

In any body of water in which oxygen-rich surface waters are well-mixed (holomictic), no chemocline will exist, as there is no stratification of layers. [7] Chemoclines can become unstable when dissolved gases become supersaturated, such as H2S, due to mixing associated with bubbling or boiling (ebullition). [8]

Chemocline structure

Position of the chemocline between oxic and anoxic layers Chemoclines.png
Position of the chemocline between oxic and anoxic layers
Purple bacteria pumped from a chemocline 7 meter deep in a meromictic lake Chemocline water Lake Mahoney.jpg
Purple bacteria pumped from a chemocline 7 meter deep in a meromictic lake

Containing the largest chemical gradient, the chemocline is a thin boundary layer that separates a meromictic lake into two parts: the upper mixolimnion and the lower monimolimnion. [7] The mixolimnion is a region that is accessed by the wind where the water can be fully mixed and circulated. However, the monimolimnion is dense and cannot interact with the wind in the same manner, preventing mixing. Furthermore, the chemocline's variability in density determines the degree to which the body of water will experience mixing and circulation. Since the chemocline acts as a barrier between the mixed and non-mixed layers, the deeper monimolimnion layer is often anoxic. [1] A lack of gas exchange between the monimolimnion layer and the atmosphere causes an increase in oxygen consumption over oxygen production. This creates a negative redox potential along with anoxic and euxinia conditions. [7]

Chemocline instability is characterized by vertical mixing events. These can be triggered by an increase in H2S concentrations higher than 1 mmol/kg in the sulfide-rich deep monolimnion layer. The euxinic deep water would then upwell into the mixolimnion near the surface and hydrogen sulfide would be expelled into the atmosphere. [8] This can also be triggered by other gases such as carbon dioxide.

In many lakes, chemocline instability is typical. Lake stratification can be upset due to mixing events that occur 1, 2, or more times per year. These mixing events occur in monomictic, dimictic, or polymictic lakes. However, in meromictic lakes, stratification is permanent. These lakes, with a stable chemocline, are typically narrow and deep with low surface to volume ratios, low wind disturbance, and ongoing eutrophication. [6]

Life and chemoclines

As a result of the differences between the upper and lower layers, aerobic life is restricted to the region above the chemocline, while anaerobic species able to live in anoxic conditions reside below the cline. Additionally, above the chemocline, photosynthetic processes can occur due to the presence of light, but below, sufficient light is not present for photosynthetic bacteria to thrive. [9] In the mixolimnion, above the chemocline, examples of phototrophic species include cyanobacteria, while the monolimnion contains sulfate reducers and sulfide oxidizers. [7] At the chemocline itself, photosynthetic forms of anaerobic bacteria, like green phototrophic and purple sulfur bacteria, cluster and take advantage of both the sunlight from above and the hydrogen sulfide (H2S) produced by the anaerobic bacteria below. [7] [9] Due to the gradient of conditions, the chemocline layer may contain an abundance of phototrophic bacteria and high concentrations of thiosulfate and elemental sulfur. [7] Methanotrophic bacteria have also been found in the anoxic gradient of some chemoclines. [10] A study conducted in Ace Lake, located in Antarctica, investigated the process of anoxygenic photosynthesis done by green sulfur bacteria in the lake and found that they were located exclusively in the chemocline of the lake due to the presence of light and sulfide. [9]

Furthermore, microbial processes can be responsible for the presence of chemical differences in a chemocline. Processes like carbon dioxide fixation, sulfur cycling, and exoenzyme activities occur at heightened rates in the cline compared to the surrounding body of water. Because of the various chemical properties of a chemocline, it can often support a diverse array of lifeforms in a small layer. [11]

However, chemocline instability can upset the balance of bacterial species found in each layer. Euxinic deep water that upwells into the photic zone can introduce sulfides and cause a bloom of sulfur oxidizing bacteria in the upper mixolimnion. [8]

Related Research Articles

<span class="mw-page-title-main">Green sulfur bacteria</span> Family of bacteria

The green sulfur bacteria are a phylum, Chlorobiota, of obligately anaerobic photoautotrophic bacteria that metabolize sulfur.

The purple sulfur bacteria (PSB) are part of a group of Pseudomonadota capable of photosynthesis, collectively referred to as purple bacteria. They are anaerobic or microaerophilic, and are often found in stratified water environments including hot springs, stagnant water bodies, as well as microbial mats in intertidal zones. Unlike plants, algae, and cyanobacteria, purple sulfur bacteria do not use water as their reducing agent, and therefore do not produce oxygen. Instead, they can use sulfur in the form of sulfide, or thiosulfate (as well, some species can use H2, Fe2+, or NO2) as the electron donor in their photosynthetic pathways. The sulfur is oxidized to produce granules of elemental sulfur. This, in turn, may be oxidized to form sulfuric acid.

<span class="mw-page-title-main">Meromictic lake</span> Permanently stratified lake with layers of water that do not intermix

A meromictic lake is a lake which has layers of water that do not intermix. In ordinary, holomictic lakes, at least once each year, there is a physical mixing of the surface and the deep waters.

<span class="mw-page-title-main">Purple bacteria</span> Group of phototrophic bacteria

Purple bacteria or purple photosynthetic bacteria are Gram-negative proteobacteria that are phototrophic, capable of producing their own food via photosynthesis. They are pigmented with bacteriochlorophyll a or b, together with various carotenoids, which give them colours ranging between purple, red, brown, and orange. They may be divided into two groups – purple sulfur bacteria and purple non-sulfur bacteria. Purple bacteria are anoxygenic phototrophs widely spread in nature, but especially in aquatic environments, where there are anoxic conditions that favor the synthesis of their pigments.

<span class="mw-page-title-main">Green Lake (New York)</span> Lake in New York

Green Lake is the larger of the two lakes in Green Lakes State Park, which lies about 9 miles (14 km) east of downtown Syracuse in Onondaga County, New York. Round Lake is the smaller lake located west of Green Lake. Both lakes are meromictic, which means no seasonal mixing of surface and bottom waters occurs. Meromictic lakes are fairly rare; they have been extensively studied, in part because their sediments can preserve a historical record extending back thousands of years, and because of the euxinic conditions which can form in the deep water.

<span class="mw-page-title-main">Chromatiaceae</span> Family of purple sulfur bacteria

The Chromatiaceae are one of the two families of purple sulfur bacteria, together with the Ectothiorhodospiraceae. They belong to the order Chromatiales of the class Gammaproteobacteria, which is composed by unicellular Gram-negative organisms. Most of the species are photolithoautotrophs and conduct an anoxygenic photosynthesis, but there are also representatives capable of growing under dark and/or microaerobic conditions as either chemolithoautotrophs or chemoorganoheterotrophs.

<i>Beggiatoa</i> Genus of bacteria

Beggiatoa is a genus of Gammaproteobacteria belonging to the order Thiotrichales, in the Pseudomonadota phylum. These bacteria form colorless filaments composed of cells that can be up to 200 µm in diameter, and are one of the largest prokaryotes on Earth. Beggiatoa are chemolithotrophic sulfur-oxidizers, using reduced sulfur species as an energy source. They live in sulfur-rich environments such as soil, both marine and freshwater, in the deep sea hydrothermal vents, and in polluted marine environments. In association with other sulfur bacteria, e.g. Thiothrix, they can form biofilms that are visible to the naked eye as mats of long white filaments; the white color is due to sulfur globules stored inside the cells.

<span class="mw-page-title-main">Winogradsky column</span> Device for culturing microorganisms

The Winogradsky column is a simple device for culturing a large diversity of microorganisms. Invented in the 1880s by Sergei Winogradsky, the device is a column of pond mud and water mixed with a carbon source such as newspaper, blackened marshmallows or egg-shells, and a sulfur source such as gypsum or egg yolk. Incubating the column in sunlight for months results in an aerobic/anaerobic gradient as well as a sulfide gradient. These two gradients promote the growth of different microorganisms such as Clostridium, Desulfovibrio, Chlorobium, Chromatium, Rhodomicrobium, and Beggiatoa, as well as many other species of bacteria, cyanobacteria, and algae.

Anoxic waters are areas of sea water, fresh water, or groundwater that are depleted of dissolved oxygen. The US Geological Survey defines anoxic groundwater as those with dissolved oxygen concentration of less than 0.5 milligrams per litre. Anoxic waters can be contrasted with hypoxic waters, which are low in dissolved oxygen. This condition is generally found in areas that have restricted water exchange.

Microbial metabolism is the means by which a microbe obtains the energy and nutrients it needs to live and reproduce. Microbes use many different types of metabolic strategies and species can often be differentiated from each other based on metabolic characteristics. The specific metabolic properties of a microbe are the major factors in determining that microbe's ecological niche, and often allow for that microbe to be useful in industrial processes or responsible for biogeochemical cycles.

γ-Carotene (gamma-carotene) is a carotenoid, and is a biosynthetic intermediate for cyclized carotenoid synthesis in plants. It is formed from cyclization of lycopene by lycopene cyclase epsilon. Along with several other carotenoids, γ-carotene is a vitamer of vitamin A in herbivores and omnivores. Carotenoids with a cyclized, beta-ionone ring can be converted to vitamin A, also known as retinol, by the enzyme beta-carotene 15,15'-dioxygenase; however, the bioconversion of γ-carotene to retinol has not been well-characterized. γ-Carotene has tentatively been identified as a biomarker for green and purple sulfur bacteria in a sample from the 1.640 ± 0.003-Gyr-old Barney Creek Formation in Northern Australia which comprises marine sediments. Tentative discovery of γ-carotene in marine sediments implies a past euxinic environment, where water columns were anoxic and sulfidic. This is significant for reconstructing past oceanic conditions, but so far γ-carotene has only been potentially identified in the one measured sample.

Uet era Ngermeuangel is a marine lake located on Koror island in Palau. There are about 70 other marine lakes located throughout the Rock Islands and Koror. Uet era Ngermeuangel is notable for endemic subspecies of golden jellyfish and is one of five marine lakes in Palau used for several scientific researches in evolutionary biology the other lakes being Jellyfish Lake, Clear Lake (Palau), Goby Lake, Uet era Ongael.

<span class="mw-page-title-main">Redox gradient</span>

A redox gradient is a series of reduction-oxidation (redox) reactions sorted according to redox potential. The redox ladder displays the order in which redox reactions occur based on the free energy gained from redox pairs. These redox gradients form both spatially and temporally as a result of differences in microbial processes, chemical composition of the environment, and oxidative potential. Common environments where redox gradients exist are coastal marshes, lakes, contaminant plumes, and soils.

<span class="mw-page-title-main">Soda lake</span> Lake that is strongly alkaline

A soda lake or alkaline lake is a lake on the strongly alkaline side of neutrality, typically with a pH value between 9 and 12. They are characterized by high concentrations of carbonate salts, typically sodium carbonate, giving rise to their alkalinity. In addition, many soda lakes also contain high concentrations of sodium chloride and other dissolved salts, making them saline or hypersaline lakes as well. High pH and salinity often coincide, because of how soda lakes develop. The resulting hypersaline and highly alkalic soda lakes are considered some of the most extreme aquatic environments on Earth.

<span class="mw-page-title-main">Isorenieratene</span> Chemical compound

Isorenieratene /ˌaɪsoʊrəˈnɪərətiːn/ is a carotenoid light harvesting pigment produced exclusively by the genus Chlorobium. Chlorobium are the brown-colored strains of the family of green sulfur bacteria (Chlorobiaceae). Green sulfur bacteria are anaerobic photoautotrophic organisms meaning they perform photosynthesis in the absence of oxygen using hydrogen sulfide in the following reaction:

Chlorobium chlorochromatii, originally known as Chlorobium aggregatum, is a symbiotic green sulfur bacteria that performs anoxygenic photosynthesis and functions as an obligate photoautotroph using reduced sulfur species as electron donors. Chlorobium chlorochromatii can be found in stratified freshwater lakes.

Okenane, the diagenetic end product of okenone, is a biomarker for Chromatiaceae, the purple sulfur bacteria. These anoxygenic phototrophs use light for energy and sulfide as their electron donor and sulfur source. Discovery of okenane in marine sediments implies a past euxinic environment, where water columns were anoxic and sulfidic. This is potentially tremendously important for reconstructing past oceanic conditions, but so far okenane has only been identified in one Paleoproterozoic rock sample from Northern Australia.

Euxinia or euxinic conditions occur when water is both anoxic and sulfidic. This means that there is no oxygen (O2) and a raised level of free hydrogen sulfide (H2S). Euxinic bodies of water are frequently strongly stratified; have an oxic, highly productive, thin surface layer; and have anoxic, sulfidic bottom water. The word "euxinia" is derived from the Greek name for the Black Sea (Εὔξεινος Πόντος (Euxeinos Pontos)) which translates to "hospitable sea". Euxinic deep water is a key component of the Canfield ocean, a model of oceans during the Proterozoic period (known as the Boring Billion) proposed by Donald Canfield, an American geologist, in 1998. There is still debate within the scientific community on both the duration and frequency of euxinic conditions in the ancient oceans. Euxinia is relatively rare in modern bodies of water, but does still happen in places like the Black Sea and certain fjords.

Chlorobactane is the diagenetic product of an aromatic carotenoid produced uniquely by green-pigmented green sulfur bacteria (GSB) in the order Chlorobiales. Observed in organic matter as far back as the Paleoproterozoic, its identity as a diagnostic biomarker has been used to interpret ancient environments.

Thiodictyon is a genus of gram-negative bacterium classified within purple sulfur bacteria (PSB).

References

  1. 1 2 3 Stewart KM, Walker KF, Likens GE (2009). "Meromictic Lakes". Encyclopedia of Inland Waters. Elsevier. pp. 589–602. doi:10.1016/b978-012370626-3.00027-2. ISBN   9780123706263.
  2. Uveges BT, Junium CK, Scholz CA, Fulton JM (2020-10-15). "Chemocline collapse in Lake Kivu as an analogue for nitrogen cycling during Oceanic Anoxic Events". Earth and Planetary Science Letters. 548: 116459. Bibcode:2020E&PSL.54816459U. doi: 10.1016/j.epsl.2020.116459 . ISSN   0012-821X. S2CID   224981010.
  3. Danza F, Storelli N, Roman S, Lüdin S, Tonolla M (2017-12-15). "Dynamic cellular complexity of anoxygenic phototrophic sulfur bacteria in the chemocline of meromictic Lake Cadagno". PLOS ONE. 12 (12): e0189510. Bibcode:2017PLoSO..1289510D. doi: 10.1371/journal.pone.0189510 . PMC   5731995 . PMID   29245157.
  4. Sinninghe Damsté JS, de Leeuw JW, Wakeham SG, Hayes JM, Kohnen ME (1993-12-02). "Chemocline of the Black Sea". Nature. 366 (6454): 416. Bibcode:1993Natur.366..416S. doi: 10.1038/366416a0 . ISSN   1476-4687. S2CID   11974369.
  5. Oikonomou A, Filker S, Breiner HW, Stoeck T (June 2015). "Protistan diversity in a permanently stratified meromictic lake (Lake Alatsee, SW Germany)". Environmental Microbiology. 17 (6): 2144–2157. Bibcode:2015EnvMi..17.2144O. doi:10.1111/1462-2920.12666. PMID   25330396.
  6. 1 2 Blees J, Niemann H, Wenk CB, Zopfi J, Schubert CJ, Kirf MK, et al. (2014-01-27). "Micro-aerobic bacterial methane oxidation in the chemocline and anoxic water column of deep south-Alpine Lake Lugano (Switzerland)". Limnology and Oceanography. 59 (2): 311–324. Bibcode:2014LimOc..59..311B. doi: 10.4319/lo.2014.59.2.0311 . ISSN   0024-3590. S2CID   56401767.
  7. 1 2 3 4 5 6 Čanković, M; Žućko, J; Petrić, I; Marguš, M; Ciglenecćki, I (2020-05-14). "Impact of euxinic holomictic conditions on prokaryotic assemblages in a marine meromictic lake". Aquatic Microbial Ecology. 84: 141–154. doi:10.3354/ame01931. ISSN   0948-3055. S2CID   216313230.
  8. 1 2 3 Riccardi, Anthony L.; Arthur, Michael A.; Kump, Lee R. (2006-12-01). "Sulfur isotopic evidence for chemocline upward excursions during the end-Permian mass extinction". Geochimica et Cosmochimica Acta. A Special Issue Dedicated to Robert A. Berner. 70 (23): 5740–5752. Bibcode:2006GeCoA..70.5740R. doi:10.1016/j.gca.2006.08.005. ISSN   0016-7037.
  9. 1 2 3 Neretin LE (2006). Past and present water column anoxia. Dordrecht: Springer. ISBN   978-1-4020-4297-3. OCLC   209932741.
  10. Blees, Jan; Niemann, Helge; Wenk, Christine B.; Zopfi, Jakob; Schubert, Carsten J.; Kirf, Mathias K.; Veronesi, Mauro L.; Hitz, Carmen; Lehmann, Moritz F. (2014). "Micro-aerobic bacterial methane oxidation in the chemocline and anoxic water column of deep south-Alpine Lake Lugano (Switzerland)". Limnology and Oceanography. 59 (2): 311–324. Bibcode:2014LimOc..59..311B. doi: 10.4319/lo.2014.59.2.0311 . ISSN   0024-3590. S2CID   56401767.
  11. Sass AM, Sass H, Coolen MJ, Cypionka H, Overmann J (December 2001). "Microbial communities in the chemocline of a hypersaline deep-sea basin (Urania basin, Mediterranean Sea)". Applied and Environmental Microbiology. 67 (12): 5392–5402. Bibcode:2001ApEnM..67.5392S. doi:10.1128/AEM.67.12.5392-5402.2001. PMC   93321 . PMID   11722884.