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Soda lake Information

This page is about soda lakes in general. For lakes named Soda, see Soda Lake (disambiguation).
Lake Shala, in the East African Rift Valley

A soda lake or alkaline lake is a highly alkaline lake due to its high concentrations of carbonate salts, typically sodium carbonate and related salt complexes, and with a pH value over 8.5 to 9. Many soda lakes also contain high concentrations of sodium chloride and other dissolved salts, making them saline or hypersaline lakes.

They are among the most extreme aquatic environments on Earth. [1] However, despite their apparent inhospitability, they are often highly productive ecosystems, compared to their (pH-neutral) freshwater counterparts, and the most productive aquatic environments on Earth. An important cause for their high productivity is the abundance of dissolved carbon dioxide.

They are considered as environments that conserve and/or mimic ancient life conditions.

Soda lakes occur naturally throughout the world (see table below). Many are in arid and semi-arid areas and most are on the margins of tectonic plates, [2] in connection to tectonic rifts like the East African Rift Valley and other such places. [3] In Brazil, the Pantanal has around 600 shallow soda lakes. [4]

Geochemistry, geology and genesis

They are considered as environments that conserve and/or mimic ancient life conditions. [5]

topography : endorheic lake

high alkalinity

relative deficit in soluble magnesium or calcium, so that they do not alter the suaturation of carbonate ions.

Geology

Soda lakes seem to be associated with active tectonic and volcanic zones and exist in two types:

Endorheic lake
One condition is an 
endorheic basin, corresponding to a water body with no outflow of water.

There are exceptions to the "no outflow" rule: both Lake Kivu and Lake Tanganyika have outlets but also have the characteristics of soda lakes, and Lake Tanganyika even grows microbialites. [8]

Geochemistry

Alkalinity

The basic condition of a soda lake is that the total alkalinity (or TA) — that is, the amount of carbonates and bicarbonates, commonly measured in milliequivalents (of dissolved carbonates) per liter (meq/l) — is superior to the alkaline earth ions magnesium (Mg) and calcium (Ca): [9]
TA > ( Mg + Ca)
or, in more detailed form: [10]
∑([HCO3-
carbonate + 2[CO32-
bicarbonate) > ∑(2[Mg2+ion
magnesium 2[Ca2+ion
calcium)

This ratio is most likely to occur in areas with fresh volcanic rocks, where carbon dioxide (CO2) can react with fresh silicates, mobilizing sodium (Na), potassium (K), magnesium (Mg) and calcium (Ca) in the appropriate proportions. But alkalinity can also rise by sulfate reduction [11] (see " Sulfur cycle" below).

When such water is evaporated, [CO32-] (bicarbonate) gets more concentrated; this causes a rise in pH, which in turn induces Na carbonates to precipitate. [3] 

The high 
alkalinity and 
salinity arise through evaporation of the lake water. This requires suitable climatic conditions, in order for the inflow to balance the loss of water through 
evaporation. The rate at which carbonate salts are dissolved into the lake water also depends on the surrounding geology and can in some cases lead to relatively high alkalinity even in lakes with significant outflow.

Another condition for the formation of a soda lake is the relative absence of soluble 
magnesium or 
calcium. Otherwise, dissolved magnesium (Mg2+) or calcium (Ca2+) will quickly remove the carbonate ions, through the precipitation of minerals such as 
calcite, 
magnesite or 
dolomite, effectively neutralizing the pH of the lake water. This results in a neutral (or slightly basic) 
salt lake instead. A good example is the 
Dead Sea, which is very rich in Mg2+. In some soda lakes, inflow of Ca2+ through subterranean seeps, can lead to localized precipitation. In 
Mono Lake, California and 
Lake Van, Turkey, such precipitation has formed columns of 
tufa rising above the lake surface.

Ongoing weathering within the crater lakes and their hydrothermal system (as in the case of Niuafo‘ou), or continuing evaporation in endorheic lakes will then lead to mature soda lake chemistry and to a CaCO3 super-saturation (saturation index or SI = 1) that can sustain 
microbialite growth.
[11]

Therefore a lake can become alkalic if particular geographical, geological and climatic conditions are combined.
Stratification
Tufa columns at Mono Lake, California

Many soda lakes are strongly stratified, with a well-oxygenated upper layer ( epilimnion) and an anoxic lower layer ( hypolimnion), without oxygen and often high concentrations of sulfide. Stratification can be permanent, or with seasonal mixing. The depth of the oxic/anoxic interface separating the two layers varies from a few centimeters to near the bottom sediments, depending on local conditions. In either case, it represents an important barrier, both physically and between strongly contrasting biochemical conditions.

Biodiversity

Soda lakes are unusually highly productive ecosystems, compared to their (pH-neutral) freshwater counterparts. [1] Gross primary production ( photosynthesis) rates above 10 g C m−2 day−1 (grams of carbon per square meter per day), over 16 times the global average for lakes and streams (0.6 g C m−2 day−1), have been measured. [12] This makes them one of the most productive aquatic environments on Earth. [13]

In the soda lake Nhecolândia (Pantanal, Brazil) and in a deep sea environment ( Campos dos Goytacazes, Brazilian Atlantic Ocean)
Mimiviridae members that surprisingly harbored a long, thick tail as they grew on Acanthamoeba castellanii and Vermamoeba vermiformis. We named these strains Tupanvirus soda lake and Tupanvirus deep ocean
This tail is the longest described in the virosphere
these giant viruses present the largest translational apparatus within the known virosphere

[5]

Micro-organisms

Contrary to freshwater ecosystems, their living organisms are often completely dominated by prokaryotes, i.e. bacteria and archaea, particularly in lakes with more "extreme" conditions (higher alkalinity and salinity or lower oxygen content). [14]

Their microbial richness and activity are also very different from that of other high-salt systems. This is essentially due to the main physico-chemical features of two dominant salts: sodium chloride (NaCl) in neutral saline systems and sodium carbonates in highly alkaline soda lakes, that influence the amount of energy required for osmotic migration of molecules. [15]

Microbial diversity

Soda lakes are inhabited by a rich diversity of microbial life, [a] often in dense concentrations. This leads to permanent or seasonal "algae blooms" with visible colouration in many lakes. The colour varies between particular lakes, depending on their predominant life forms and can range from green to orange or red. [1]

Lesser flamingos (Phoenicopterus minor) feeding on cyanobacteria in Lake Nakuru, Kenya

In general, the microbial biodiversity of soda lakes is relatively poorly studied. Many studies have focused on the primary producers, namely the photosynthesizing cyanobacteria or eukaryotic algae (see Carbon cycle). As studies have traditionally relied on microscopy, identification has been hindered by the fact that many soda lakes harbour species that are unique to these relatively unusual habitats and in many cases thought to be endemic, i.e. existing only in one lake. [17] The morphology (appearance) of algae and other organisms may also vary from lake to lake, depending on local conditions, making their identification more difficult, which has probably led to several instances of taxonomic confusions in the scientific literature.

Molecular methods such as DNA fingerprinting or sequencing have been used to study the diversity of organisms in soda lakes. [17] [18] [19] [20] [21] [b] For instance, 16S ribosomal RNA gene has revealed that the bacterial community of the lake with the highest salinity was characterized by a higher recent accelerated diversification than the community of a freshwater lake, whereas the phylogenetic diversity in the hypersaline lake was lower than that in a freshwater lake. [16]

Biogeography and uniqueness (endemism)

In addition to their rich biodiversity, soda lakes often harbour many unique species, adapted to alkalic conditions and unable to live in environments with neutral pH. These are called alkaliphiles. Among alkaliphiles organisms, those also adapted to high salinity are called haloalkaliphiles. Culture-independent genetic surveys have shown that soda lakes contain an unusually high amount of alkaliphilic microorganisms with low genetic similarity to known species. [18] [19] [20] [21] This indicates a long evolutionary history of adaptation to these habitats with few new species from other environments becoming adapted over time.

In-depth genetic surveys also show an unusually low overlap between the microbial communities present in the various soda lakes with only slightly different conditions such as pH and salinity. [14] [20] This trend is especially strong in the bottom layer ( hypolimnion) of stratified lakes, [17] probably because of the isolated character of such environments. Diversity data from soda lakes suggest the existence of many endemic microbial species, unique to individual lakes. [14] [20] This is a controversial finding, since conventional wisdom in microbial ecology dictates that most microbial species are cosmopolitan and dispersed globally, thanks to their enormous population sizes, a famous hypothesis first formulated by Lourens Baas Becking in 1934 ("Everything is everywhere, but the environment selects"). [24]

Macro-organisms

A rich diversity of eukaryotic algae, protists and fungi have also been encountered in many soda lakes. [14]

Multicellular animals such as crustaceans (notably the brine shrimp Artemia and the copepod Paradiaptomus africanus) and fish (e.g. Alcolapia), are also found in many of the less extreme soda lakes, adapted to the conditions of these alkalic and often saline environments. Particularly in the East African Rift Valley, microorganisms in soda lakes also provide the main food source for vast flocks of the lesser flamingo (Phoeniconaias minor). The cyanobacteria of the genus Arthrospira (formerly Spirulina) are a particularly preferred food source for these birds, owing to their large cell size and high nutritional value. Declines in East African soda lake productivity due to rising water levels threaten this food source. This may force lesser flamingos to move north and south, away from the equator. [13]

Chemical cycles and linked organisms

Carbon cycle, photosynthesis and methanogenesis

Cyanobacteria of the genus Arthrospira (synonymous to "Spirulina")

a combination of high phytoplankton standing crop and above-average biomass-specific rates, partly due the large reserve of CO2 for localized photosynthetic activity [25]

Photosynthesis produces chemical energy stored in intracellular organic compounds containing carbon. It dominates the activity at the surface of soda lakes and this process provides the primary energy source for life in the lake. [26]
The most important photosynthesizers are typically cyanobacteria, but in many less "extreme" soda lakes, eukaryotes such as green algae (Chlorophyta) can also dominate. The major genera of cyanobacteria typically found in soda lakes include Arthrospira (formerly Spirulina) (notably A. platensis), Anabaenopsis, [26] Cyanospira, Synechococcus or Chroococcus. [27] In more saline soda lakes, haloalkaliphilic archaea such as Halobacteria and bacteria such as Halorhodospira dominate photosynthesis. However, it is not clear whether this is an autotrophic process or if these require organic carbon from cyanobacterial blooms, occurring during periods of heavy rainfall that dilute the surface waters. [1]

Below the surface, anoxygenic photosynthesizers using other substances than carbon dioxide for photosynthesis also contribute to primary production in many soda lakes. These include purple sulfur bacteria such as Ectothiorhodospiraceae and purple non-sulfur bacteria such as Rhodobacteraceae (for example the species Rhodobaca bogoriensis isolated from Lake Bogoria [28]).

The photosynthesizing bacteria provide a food source for a vast diversity of aerobic and anaerobic organotrophic microorganisms from phyla including Pseudomonadota, Bacteroidota, Spirochaetota, Bacillota, Thermotogota, Deinococcota, Planctomycetota, Actinomycetota, Gemmatimonadota, and more. [1] [14] The anaerobic fermentation of organic compounds originating from the primary producers, results in one-carbon (C1) compounds such as methanol and methylamine.

At the bottom of lakes (in the sediment or hypolimnion), methanogens use these compounds to derive energy, by producing methane, a procedure known as methanogenesis. A diversity of methanogens including the archaeal genera Methanocalculus, Methanolobus, Methanosaeta, Methanosalsus and Methanoculleus have been found in soda lake sediments. [1] [29] When the resulting methane reaches the aerobic water of a soda lake, it can be consumed by methane-oxidizing bacteria such as Methylobacter or Methylomicrobium. [1]

Sulfur cycle

Sulfur-reducing bacteria are common in anoxic layers of soda lakes. These reduce sulfate and organic sulfur from dead cells into sulfide (S2−). Anoxic layers of soda lakes are therefore often rich in sulfide. As opposed to neutral lakes, the high pH prohibits the release of hydrogen sulfide (H2S) in gas form. Genera of alkaliphilic sulfur-reducers found in soda lakes include Desulfonatronovibrio and Desulfonatronum. [1] These also play important an ecological role besides in the cycling of sulfur, as they also consume hydrogen, resulting from the fermentation of organic matter.

Sulfur-oxidating bacteria instead derive their energy from oxidation of the sulfide reaching the oxygenated layers of soda lakes. Some of these are photosynthetic sulfur phototrophs, which means that they also require light to derive energy. Examples of alkaliphilic sulfur-oxidizing bacteria are the genera Thioalkalivibrio, Thiorhodospira, Thioalkalimicrobium and Natronhydrogenobacter. [1]

Nitrogen and other nutrients

Nitrogen is a limiting nutrient for growth in many soda lakes, making the internal nitrogen cycle very important for their ecological functioning. [30] One possible source of bio-available nitrogen is diazotrophic cyanobacteria, which can fix nitrogen from the atmosphere during photosynthesis. However, many of the dominant cyanobacteria found in soda lakes such as Arthrospira are probably not able to fix nitrogen. [1] Ammonia, a nitrogen-containing waste product from degradation of dead cells, can be lost from soda lakes through volatilization because of the high pH. This can hinder nitrification, in which ammonia is "recycled" to the bio-available form nitrate. Nevertheless, ammonia oxidation seems to be efficiently carried out in soda lakes in either case, probably by ammonia-oxidizing bacteria as well as Thaumarchaea. [30]

List of soda lakes

Pangong lake, India & Tibet
Astronaut photograph of the mostly dry bed of Owens Lake, Ca, USA
2010 satellite image of Sambhar Salt Lake, India
Lake Eyasi, Tanzania
Flamingos feeding at Lake Nakuru, Kenya
Lake Turkana, Kenya
Satonda Island lake, Indonesia
Niuafo'ou lake, Tonga
Lake Specchio di Venere, Pantelleria island, Italy

The following table lists some examples of soda lakes by region, listing country, pH and salinity. NA indicates 'data not available':

Continent Name Country pH Salinity
Africa Lake Ngami [31] Botswana
Sua Pan Botswana 19%
Rombou Lake Chad 10.2 [32] 2% [31]
Wadi El Natrun lakes Egypt 9.5 5%
Lake Arenguadi (Green Lake) Ethiopia 9.5–9.9 [14] 0.25% [31]
Lake Basaka Ethiopia 9.6 [14] 0.3% [31]
Lake Shala Ethiopia 9.8 [14] 1.8% [31]
Lake Abijatta Ethiopia 9.9 [14] 3.4% [33]
Lake Chitu [c] Ethiopia 10.3 [14] 5.8% [33]
Lake Bogoria Kenya 10.5 3.5% [33]
Empakai Crater lake [6] Kenya
Lake Logipi Kenya 9.5–10.5 2–5% [33]
Lake Magadi Kenya 10 >10% [33]
Lake Nakuru Kenya 10.5 [33] NA
Lake Sonachi (Crater Lake) [33] Kenya NA NA
Lake Turkana Kenya 8.5–9.2 [34] 0.25% [33]
Malha Crater Lake Sudan 9.5–10.3 [33] NA
Lake Balangida [33] Tanzania NA NA
Lake Eyasi Tanzania 9.3 [35] 0.5% [33]
Lake Manyara Tanzania 9.5–10 [33] [35] NA
Momela Lakes Tanzania 9.7 22%
Lake Natron Tanzania 9–10.5 >10% [33]
Lake Rukwa Tanzania 8–9 [35] [33] NA
Asia Guozha lake [33] China NA NA
Qinghai Lake China 9.3 [36] 2.2% [33]
Lake Zabuye (Drangyer) China 10 [33] NA
Kartsakhi Lake Georgia/ Turkey NA 0.09%
Khyagar Lake [32] India 9.5 0.6% [33]
Kushul lake India NA NA
Lonar Lake (Crater Lake) India 9.5–10.5 [18] 1% [33]
Namucuo Lake India 9.4 [36] 0.2% [33]
Sambhar Salt Lake India 9.5 7% [33]
Tso Kar Salt Lake India 8.8 [33] NA
Tso Moriri Salt Lake India 9.0 [33] NA
Aksayqin Hu Lake Aksai Chin, India/China NA NA [33] [37]
Lake Hongshan Hu [33] Aksai Chin, India/China NA NA
Pangong Lake India & China 9.4 0.9% [33] [38]
Spanggur Tso (Pongur Tso) [33] India & China NA NA
Surigh Yilganing Kol [33] Aksai Chin, India/China NA NA
Tianshuihai lake [33] Aksai Chin, India/China NA NA
North Tianshuihai lake Aksai Chin, India/China NA NA
Tso Tang Lake [33] Aksai Chin, India/China NA NA
Satonda Island Indonesia 8.55
Kulunda Steppe lakes ( Borli) Kazakhstan 8.89-9.16 5.7% [33] [39]
Kulunda St. ( Uyaly) Kazakhstan 9.47-9.50 2.7% [39]
Taboos-nor [33] Mongolia NA NA
Lake Khatyn Russia 10 [33] NA
Torey Lakes Russia, Mongolia NA NA
Lake Salda [33] Turkey NA NA
Lake Van Turkey 9.7–9.8 2.3% [33]
Europe Lake Neusiedl (Fertő) Austria, Hungary 9–9.3 [33] [40] NA
Böddi-szék Hungary 8.8–9.8 [33] [40] 12.34% [41]
Lake Fehér (Szeged) [33] Hungary NA NA
Kelemen-szék Hungary 9–9.7 [33] [40] [42] NA
Nagy-Vadas [42] Hungary NA NA
Specchio di Venere [6] [43] ( Pantelleria Island) Italy
Velika Rusanda [44] Serbia 9.3 [33] [40] NA
Malham Tarn UK 8.0–8.6 [45] [46] NA
North America Manitou Lake, [33] SK Canada NA NA
Deer Lake [47] ( Cariboo Plateau, BC) Canada
Goodenough Lake [47] ( Bonaparte Plateau, BC) Canada 10.2 [33] NA
Last Chance Lake [47] ( Bonaparte Plateau, BC) Canada
Probe Lake [47] ( Cariboo Plateau, BC) Canada
Lake Texcoco Mexico 8.8–11.5 8% [33]
Lake Alchichica Mexico 8.9 [33] NA
Alkali Lake, OR US 11 [33] NA
Baldwin Lake, [33] Ca US NA NA
Borax Lake, OR US NA NA
Kauhako Crater Lake [9], Molokai, HI US
Mono Lake, Ca US 9.8 [30] 8% [33]
Owens Lake, Ca [33] US NA NA
Soap Lake, WA US 9.7 0.7% [33]
Soda Lakes, NV US 9.7 [33] NA
Summer Lake, OR [33] US NA NA
South America Antofagasta Lake [33] Chile NA NA
Oceania Niuafoʻou Caldera Lake [48] Tonga
Lake Werowrap [32] Australia 9.8 4% [33]

Industrial use

Many water-soluble chemicals are extracted from soda lake waters worldwide. Lithium carbonate (see Lake Zabuye), potash (see lake Lop Nur and Qinghai Salt Lake Potash), soda ash (see Lake Abijatta and Lake Natron), etc. are extracted in large quantities. Lithium carbonate is a raw material in production of lithium which has applications in lithium storage batteries widely used in modern electronic gadgets and electrically powered automobiles. Water of some soda lakes are rich in dissolved uranium carbonate. [49] Algaculture is carried out on a commercial scale with soda lake water. [50] [51]

See also

Notes

  1. ^ Independantly from the alkalinity factor and considering only the saline content, species richness (number of species present, or diversity) in highly saline lakes is surprisingly high, often rivaling that of freshwater ecosystems. Salinity is the primary factor regulating the diversification of bacteria and archaea, including the functional gene lineages involved in the denitrification pathway or the hydrolysis of chitin. [16]
  2. ^ These molecular methods are based on DNA extracted directly from the environment and thus do not require microorganisms to be cultured. This is a major advantage, as culturing of novel microorganisms is a laborious technique known to seriously bias the outcome of diversity studies, since only about one in a hundred organisms can be cultured using standard techniques. [22] For microorganisms, the phylogenetic marker gene small subunit (SSU) ribosomal RNA is typically targeted, due to its particular properties such as existence in all cellular organisms and ability to be used as a "molecular clock" to trace the evolutionary history of an organism. [23]
  3. ^ For Lake Chitu in Ethiopia, see also:
    • Assaye, Hirut; Belay, Amha; Desse, Gulelat Haki; Gray, David (2018). "Seasonal variation in the nutrient profile of Arthrospira fusiformis biomass harvested from an Ethiopian soda lake, Lake Chitu" (PDF). Journal of Applied Phycology. 30 (3): 1597–1606. doi: 10.1007/s10811-017-1359-0. Retrieved 2024-07-14.
    • Ogato, Tadesse; Kifle, Demeke; Fetahi, Tadesse; Sitotaw, Baye (2014). "Evaluation of growth and biomass production of Arthrospira (Spirulina) fusiformis in laboratory cultures using waters from the Ethiopian soda lakes Chitu and Shala". Journal of Applied Phycology. doi: 10.1007/s10811-014-0251-4. Retrieved 2024-07-14.

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  37. ^ Chaohai, Liu; Li, Shijie; Yafeng, Shi (January 1992). "Glacial and lake fluctuations in the area of the west Kunlun mountains during the last 45000 years". Annals of Glaciology. pp. 79–84. Retrieved 2024-07-05.
  38. ^ T.V. Ramachandra; Rao, K. Sankara; Boominathan, M.; Mahapatra, Durga Madhab; Bhat, Harish R. (February 2011). "Environmental impact assessment of the national large solar telescope project and its ecological impact in Merak area" (EIA study near Pangong Tso lake, India. CES Technical Report 123). Retrieved 2024-07-05.
  39. ^ a b Ubaskin, A; Kassanova, A; Lunkov, A; Ahmetov, K; Almagambetova, K; Erzhanov, N; Abylkhassanov, T (2020). "Hydrochemical Research and Geochemical Classification of Salt Lakes in the Pavlodar Region". IOP Conf. Ser.: Mater. Sci. Eng. (754): 012009. Retrieved 2024-07-05.
  40. ^ a b c d Felföldi, T. S.; Somogyi, B. R.; Márialigeti, K. R.; Vörös, L. (2009). "Characterization of photoautotrophic picoplankton assemblages in turbid, alkaline lakes of the Carpathian Basin (Central Europe)". Journal of Limnology. 68 (2): 385. doi: 10.4081/jlimnol.2009.385.
  41. ^ Borsodi, Andrea K; Knáb, Mónika; Czeibert, Katalin; Márialigeti, Károly; Vörös, Lajos; Somogyi, Boglárka (2013). "Planktonic bacterial community composition of an extremely shallow soda pond during a phytoplankton bloom revealed by cultivation and molecular cloning" (PDF). Extremophiles. 17 (4): 575–584. ISSN  1431-0651. Retrieved 2024-07-05.
  42. ^ a b Rusznyák, Anna; Vladar, Péter; Szabó, Gitta; Márialigeti, Károly; Borsodi, Andrea K (2008). "Bacterial diversity of reed (Phragmites australis) periphyton communities of Kelemen-szék and Nagy-Vadas (two Hungarian soda ponds)". Extremophiles. 12: 763–773. doi: 10.1007/s00792-008-0183-5.
  43. ^ Cangemi, Marianna; Censi, Paolo; Reimer, Andreas; D'Alessandro, Walter; Hause-Reitner, Dorothea; Madonia, Paolo; Oliveri, Ygor; Pecoraino, Giovannella; Reitner, Joachim (April 2016). "Carbonate precipitation in the alkaline lake Specchio di Venere (Pantelleria Island, Italy) and the possible role of microbial mats". Applied Geochemistry. 67: 168–176. Retrieved 2024-07-05.
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Bibliography

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From Wikipedia, the free encyclopedia

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Soda lake Information

This page is about soda lakes in general. For lakes named Soda, see Soda Lake (disambiguation).
Lake Shala, in the East African Rift Valley

A soda lake or alkaline lake is a highly alkaline lake due to its high concentrations of carbonate salts, typically sodium carbonate and related salt complexes, and with a pH value over 8.5 to 9. Many soda lakes also contain high concentrations of sodium chloride and other dissolved salts, making them saline or hypersaline lakes.

They are among the most extreme aquatic environments on Earth. [1] However, despite their apparent inhospitability, they are often highly productive ecosystems, compared to their (pH-neutral) freshwater counterparts, and the most productive aquatic environments on Earth. An important cause for their high productivity is the abundance of dissolved carbon dioxide.

They are considered as environments that conserve and/or mimic ancient life conditions.

Soda lakes occur naturally throughout the world (see table below). Many are in arid and semi-arid areas and most are on the margins of tectonic plates, [2] in connection to tectonic rifts like the East African Rift Valley and other such places. [3] In Brazil, the Pantanal has around 600 shallow soda lakes. [4]

Geochemistry, geology and genesis

They are considered as environments that conserve and/or mimic ancient life conditions. [5]

topography : endorheic lake

high alkalinity

relative deficit in soluble magnesium or calcium, so that they do not alter the suaturation of carbonate ions.

Geology

Soda lakes seem to be associated with active tectonic and volcanic zones and exist in two types:

Endorheic lake
One condition is an 
endorheic basin, corresponding to a water body with no outflow of water.

There are exceptions to the "no outflow" rule: both Lake Kivu and Lake Tanganyika have outlets but also have the characteristics of soda lakes, and Lake Tanganyika even grows microbialites. [8]

Geochemistry

Alkalinity

The basic condition of a soda lake is that the total alkalinity (or TA) — that is, the amount of carbonates and bicarbonates, commonly measured in milliequivalents (of dissolved carbonates) per liter (meq/l) — is superior to the alkaline earth ions magnesium (Mg) and calcium (Ca): [9]
TA > ( Mg + Ca)
or, in more detailed form: [10]
∑([HCO3-
carbonate + 2[CO32-
bicarbonate) > ∑(2[Mg2+ion
magnesium 2[Ca2+ion
calcium)

This ratio is most likely to occur in areas with fresh volcanic rocks, where carbon dioxide (CO2) can react with fresh silicates, mobilizing sodium (Na), potassium (K), magnesium (Mg) and calcium (Ca) in the appropriate proportions. But alkalinity can also rise by sulfate reduction [11] (see " Sulfur cycle" below).

When such water is evaporated, [CO32-] (bicarbonate) gets more concentrated; this causes a rise in pH, which in turn induces Na carbonates to precipitate. [3] 

The high 
alkalinity and 
salinity arise through evaporation of the lake water. This requires suitable climatic conditions, in order for the inflow to balance the loss of water through 
evaporation. The rate at which carbonate salts are dissolved into the lake water also depends on the surrounding geology and can in some cases lead to relatively high alkalinity even in lakes with significant outflow.

Another condition for the formation of a soda lake is the relative absence of soluble 
magnesium or 
calcium. Otherwise, dissolved magnesium (Mg2+) or calcium (Ca2+) will quickly remove the carbonate ions, through the precipitation of minerals such as 
calcite, 
magnesite or 
dolomite, effectively neutralizing the pH of the lake water. This results in a neutral (or slightly basic) 
salt lake instead. A good example is the 
Dead Sea, which is very rich in Mg2+. In some soda lakes, inflow of Ca2+ through subterranean seeps, can lead to localized precipitation. In 
Mono Lake, California and 
Lake Van, Turkey, such precipitation has formed columns of 
tufa rising above the lake surface.

Ongoing weathering within the crater lakes and their hydrothermal system (as in the case of Niuafo‘ou), or continuing evaporation in endorheic lakes will then lead to mature soda lake chemistry and to a CaCO3 super-saturation (saturation index or SI = 1) that can sustain 
microbialite growth.
[11]

Therefore a lake can become alkalic if particular geographical, geological and climatic conditions are combined.
Stratification
Tufa columns at Mono Lake, California

Many soda lakes are strongly stratified, with a well-oxygenated upper layer ( epilimnion) and an anoxic lower layer ( hypolimnion), without oxygen and often high concentrations of sulfide. Stratification can be permanent, or with seasonal mixing. The depth of the oxic/anoxic interface separating the two layers varies from a few centimeters to near the bottom sediments, depending on local conditions. In either case, it represents an important barrier, both physically and between strongly contrasting biochemical conditions.

Biodiversity

Soda lakes are unusually highly productive ecosystems, compared to their (pH-neutral) freshwater counterparts. [1] Gross primary production ( photosynthesis) rates above 10 g C m−2 day−1 (grams of carbon per square meter per day), over 16 times the global average for lakes and streams (0.6 g C m−2 day−1), have been measured. [12] This makes them one of the most productive aquatic environments on Earth. [13]

In the soda lake Nhecolândia (Pantanal, Brazil) and in a deep sea environment ( Campos dos Goytacazes, Brazilian Atlantic Ocean)
Mimiviridae members that surprisingly harbored a long, thick tail as they grew on Acanthamoeba castellanii and Vermamoeba vermiformis. We named these strains Tupanvirus soda lake and Tupanvirus deep ocean
This tail is the longest described in the virosphere
these giant viruses present the largest translational apparatus within the known virosphere

[5]

Micro-organisms

Contrary to freshwater ecosystems, their living organisms are often completely dominated by prokaryotes, i.e. bacteria and archaea, particularly in lakes with more "extreme" conditions (higher alkalinity and salinity or lower oxygen content). [14]

Their microbial richness and activity are also very different from that of other high-salt systems. This is essentially due to the main physico-chemical features of two dominant salts: sodium chloride (NaCl) in neutral saline systems and sodium carbonates in highly alkaline soda lakes, that influence the amount of energy required for osmotic migration of molecules. [15]

Microbial diversity

Soda lakes are inhabited by a rich diversity of microbial life, [a] often in dense concentrations. This leads to permanent or seasonal "algae blooms" with visible colouration in many lakes. The colour varies between particular lakes, depending on their predominant life forms and can range from green to orange or red. [1]

Lesser flamingos (Phoenicopterus minor) feeding on cyanobacteria in Lake Nakuru, Kenya

In general, the microbial biodiversity of soda lakes is relatively poorly studied. Many studies have focused on the primary producers, namely the photosynthesizing cyanobacteria or eukaryotic algae (see Carbon cycle). As studies have traditionally relied on microscopy, identification has been hindered by the fact that many soda lakes harbour species that are unique to these relatively unusual habitats and in many cases thought to be endemic, i.e. existing only in one lake. [17] The morphology (appearance) of algae and other organisms may also vary from lake to lake, depending on local conditions, making their identification more difficult, which has probably led to several instances of taxonomic confusions in the scientific literature.

Molecular methods such as DNA fingerprinting or sequencing have been used to study the diversity of organisms in soda lakes. [17] [18] [19] [20] [21] [b] For instance, 16S ribosomal RNA gene has revealed that the bacterial community of the lake with the highest salinity was characterized by a higher recent accelerated diversification than the community of a freshwater lake, whereas the phylogenetic diversity in the hypersaline lake was lower than that in a freshwater lake. [16]

Biogeography and uniqueness (endemism)

In addition to their rich biodiversity, soda lakes often harbour many unique species, adapted to alkalic conditions and unable to live in environments with neutral pH. These are called alkaliphiles. Among alkaliphiles organisms, those also adapted to high salinity are called haloalkaliphiles. Culture-independent genetic surveys have shown that soda lakes contain an unusually high amount of alkaliphilic microorganisms with low genetic similarity to known species. [18] [19] [20] [21] This indicates a long evolutionary history of adaptation to these habitats with few new species from other environments becoming adapted over time.

In-depth genetic surveys also show an unusually low overlap between the microbial communities present in the various soda lakes with only slightly different conditions such as pH and salinity. [14] [20] This trend is especially strong in the bottom layer ( hypolimnion) of stratified lakes, [17] probably because of the isolated character of such environments. Diversity data from soda lakes suggest the existence of many endemic microbial species, unique to individual lakes. [14] [20] This is a controversial finding, since conventional wisdom in microbial ecology dictates that most microbial species are cosmopolitan and dispersed globally, thanks to their enormous population sizes, a famous hypothesis first formulated by Lourens Baas Becking in 1934 ("Everything is everywhere, but the environment selects"). [24]

Macro-organisms

A rich diversity of eukaryotic algae, protists and fungi have also been encountered in many soda lakes. [14]

Multicellular animals such as crustaceans (notably the brine shrimp Artemia and the copepod Paradiaptomus africanus) and fish (e.g. Alcolapia), are also found in many of the less extreme soda lakes, adapted to the conditions of these alkalic and often saline environments. Particularly in the East African Rift Valley, microorganisms in soda lakes also provide the main food source for vast flocks of the lesser flamingo (Phoeniconaias minor). The cyanobacteria of the genus Arthrospira (formerly Spirulina) are a particularly preferred food source for these birds, owing to their large cell size and high nutritional value. Declines in East African soda lake productivity due to rising water levels threaten this food source. This may force lesser flamingos to move north and south, away from the equator. [13]

Chemical cycles and linked organisms

Carbon cycle, photosynthesis and methanogenesis

Cyanobacteria of the genus Arthrospira (synonymous to "Spirulina")

a combination of high phytoplankton standing crop and above-average biomass-specific rates, partly due the large reserve of CO2 for localized photosynthetic activity [25]

Photosynthesis produces chemical energy stored in intracellular organic compounds containing carbon. It dominates the activity at the surface of soda lakes and this process provides the primary energy source for life in the lake. [26]
The most important photosynthesizers are typically cyanobacteria, but in many less "extreme" soda lakes, eukaryotes such as green algae (Chlorophyta) can also dominate. The major genera of cyanobacteria typically found in soda lakes include Arthrospira (formerly Spirulina) (notably A. platensis), Anabaenopsis, [26] Cyanospira, Synechococcus or Chroococcus. [27] In more saline soda lakes, haloalkaliphilic archaea such as Halobacteria and bacteria such as Halorhodospira dominate photosynthesis. However, it is not clear whether this is an autotrophic process or if these require organic carbon from cyanobacterial blooms, occurring during periods of heavy rainfall that dilute the surface waters. [1]

Below the surface, anoxygenic photosynthesizers using other substances than carbon dioxide for photosynthesis also contribute to primary production in many soda lakes. These include purple sulfur bacteria such as Ectothiorhodospiraceae and purple non-sulfur bacteria such as Rhodobacteraceae (for example the species Rhodobaca bogoriensis isolated from Lake Bogoria [28]).

The photosynthesizing bacteria provide a food source for a vast diversity of aerobic and anaerobic organotrophic microorganisms from phyla including Pseudomonadota, Bacteroidota, Spirochaetota, Bacillota, Thermotogota, Deinococcota, Planctomycetota, Actinomycetota, Gemmatimonadota, and more. [1] [14] The anaerobic fermentation of organic compounds originating from the primary producers, results in one-carbon (C1) compounds such as methanol and methylamine.

At the bottom of lakes (in the sediment or hypolimnion), methanogens use these compounds to derive energy, by producing methane, a procedure known as methanogenesis. A diversity of methanogens including the archaeal genera Methanocalculus, Methanolobus, Methanosaeta, Methanosalsus and Methanoculleus have been found in soda lake sediments. [1] [29] When the resulting methane reaches the aerobic water of a soda lake, it can be consumed by methane-oxidizing bacteria such as Methylobacter or Methylomicrobium. [1]

Sulfur cycle

Sulfur-reducing bacteria are common in anoxic layers of soda lakes. These reduce sulfate and organic sulfur from dead cells into sulfide (S2−). Anoxic layers of soda lakes are therefore often rich in sulfide. As opposed to neutral lakes, the high pH prohibits the release of hydrogen sulfide (H2S) in gas form. Genera of alkaliphilic sulfur-reducers found in soda lakes include Desulfonatronovibrio and Desulfonatronum. [1] These also play important an ecological role besides in the cycling of sulfur, as they also consume hydrogen, resulting from the fermentation of organic matter.

Sulfur-oxidating bacteria instead derive their energy from oxidation of the sulfide reaching the oxygenated layers of soda lakes. Some of these are photosynthetic sulfur phototrophs, which means that they also require light to derive energy. Examples of alkaliphilic sulfur-oxidizing bacteria are the genera Thioalkalivibrio, Thiorhodospira, Thioalkalimicrobium and Natronhydrogenobacter. [1]

Nitrogen and other nutrients

Nitrogen is a limiting nutrient for growth in many soda lakes, making the internal nitrogen cycle very important for their ecological functioning. [30] One possible source of bio-available nitrogen is diazotrophic cyanobacteria, which can fix nitrogen from the atmosphere during photosynthesis. However, many of the dominant cyanobacteria found in soda lakes such as Arthrospira are probably not able to fix nitrogen. [1] Ammonia, a nitrogen-containing waste product from degradation of dead cells, can be lost from soda lakes through volatilization because of the high pH. This can hinder nitrification, in which ammonia is "recycled" to the bio-available form nitrate. Nevertheless, ammonia oxidation seems to be efficiently carried out in soda lakes in either case, probably by ammonia-oxidizing bacteria as well as Thaumarchaea. [30]

List of soda lakes

Pangong lake, India & Tibet
Astronaut photograph of the mostly dry bed of Owens Lake, Ca, USA
2010 satellite image of Sambhar Salt Lake, India
Lake Eyasi, Tanzania
Flamingos feeding at Lake Nakuru, Kenya
Lake Turkana, Kenya
Satonda Island lake, Indonesia
Niuafo'ou lake, Tonga
Lake Specchio di Venere, Pantelleria island, Italy

The following table lists some examples of soda lakes by region, listing country, pH and salinity. NA indicates 'data not available':

Continent Name Country pH Salinity
Africa Lake Ngami [31] Botswana
Sua Pan Botswana 19%
Rombou Lake Chad 10.2 [32] 2% [31]
Wadi El Natrun lakes Egypt 9.5 5%
Lake Arenguadi (Green Lake) Ethiopia 9.5–9.9 [14] 0.25% [31]
Lake Basaka Ethiopia 9.6 [14] 0.3% [31]
Lake Shala Ethiopia 9.8 [14] 1.8% [31]
Lake Abijatta Ethiopia 9.9 [14] 3.4% [33]
Lake Chitu [c] Ethiopia 10.3 [14] 5.8% [33]
Lake Bogoria Kenya 10.5 3.5% [33]
Empakai Crater lake [6] Kenya
Lake Logipi Kenya 9.5–10.5 2–5% [33]
Lake Magadi Kenya 10 >10% [33]
Lake Nakuru Kenya 10.5 [33] NA
Lake Sonachi (Crater Lake) [33] Kenya NA NA
Lake Turkana Kenya 8.5–9.2 [34] 0.25% [33]
Malha Crater Lake Sudan 9.5–10.3 [33] NA
Lake Balangida [33] Tanzania NA NA
Lake Eyasi Tanzania 9.3 [35] 0.5% [33]
Lake Manyara Tanzania 9.5–10 [33] [35] NA
Momela Lakes Tanzania 9.7 22%
Lake Natron Tanzania 9–10.5 >10% [33]
Lake Rukwa Tanzania 8–9 [35] [33] NA
Asia Guozha lake [33] China NA NA
Qinghai Lake China 9.3 [36] 2.2% [33]
Lake Zabuye (Drangyer) China 10 [33] NA
Kartsakhi Lake Georgia/ Turkey NA 0.09%
Khyagar Lake [32] India 9.5 0.6% [33]
Kushul lake India NA NA
Lonar Lake (Crater Lake) India 9.5–10.5 [18] 1% [33]
Namucuo Lake India 9.4 [36] 0.2% [33]
Sambhar Salt Lake India 9.5 7% [33]
Tso Kar Salt Lake India 8.8 [33] NA
Tso Moriri Salt Lake India 9.0 [33] NA
Aksayqin Hu Lake Aksai Chin, India/China NA NA [33] [37]
Lake Hongshan Hu [33] Aksai Chin, India/China NA NA
Pangong Lake India & China 9.4 0.9% [33] [38]
Spanggur Tso (Pongur Tso) [33] India & China NA NA
Surigh Yilganing Kol [33] Aksai Chin, India/China NA NA
Tianshuihai lake [33] Aksai Chin, India/China NA NA
North Tianshuihai lake Aksai Chin, India/China NA NA
Tso Tang Lake [33] Aksai Chin, India/China NA NA
Satonda Island Indonesia 8.55
Kulunda Steppe lakes ( Borli) Kazakhstan 8.89-9.16 5.7% [33] [39]
Kulunda St. ( Uyaly) Kazakhstan 9.47-9.50 2.7% [39]
Taboos-nor [33] Mongolia NA NA
Lake Khatyn Russia 10 [33] NA
Torey Lakes Russia, Mongolia NA NA
Lake Salda [33] Turkey NA NA
Lake Van Turkey 9.7–9.8 2.3% [33]
Europe Lake Neusiedl (Fertő) Austria, Hungary 9–9.3 [33] [40] NA
Böddi-szék Hungary 8.8–9.8 [33] [40] 12.34% [41]
Lake Fehér (Szeged) [33] Hungary NA NA
Kelemen-szék Hungary 9–9.7 [33] [40] [42] NA
Nagy-Vadas [42] Hungary NA NA
Specchio di Venere [6] [43] ( Pantelleria Island) Italy
Velika Rusanda [44] Serbia 9.3 [33] [40] NA
Malham Tarn UK 8.0–8.6 [45] [46] NA
North America Manitou Lake, [33] SK Canada NA NA
Deer Lake [47] ( Cariboo Plateau, BC) Canada
Goodenough Lake [47] ( Bonaparte Plateau, BC) Canada 10.2 [33] NA
Last Chance Lake [47] ( Bonaparte Plateau, BC) Canada
Probe Lake [47] ( Cariboo Plateau, BC) Canada
Lake Texcoco Mexico 8.8–11.5 8% [33]
Lake Alchichica Mexico 8.9 [33] NA
Alkali Lake, OR US 11 [33] NA
Baldwin Lake, [33] Ca US NA NA
Borax Lake, OR US NA NA
Kauhako Crater Lake [9], Molokai, HI US
Mono Lake, Ca US 9.8 [30] 8% [33]
Owens Lake, Ca [33] US NA NA
Soap Lake, WA US 9.7 0.7% [33]
Soda Lakes, NV US 9.7 [33] NA
Summer Lake, OR [33] US NA NA
South America Antofagasta Lake [33] Chile NA NA
Oceania Niuafoʻou Caldera Lake [48] Tonga
Lake Werowrap [32] Australia 9.8 4% [33]

Industrial use

Many water-soluble chemicals are extracted from soda lake waters worldwide. Lithium carbonate (see Lake Zabuye), potash (see lake Lop Nur and Qinghai Salt Lake Potash), soda ash (see Lake Abijatta and Lake Natron), etc. are extracted in large quantities. Lithium carbonate is a raw material in production of lithium which has applications in lithium storage batteries widely used in modern electronic gadgets and electrically powered automobiles. Water of some soda lakes are rich in dissolved uranium carbonate. [49] Algaculture is carried out on a commercial scale with soda lake water. [50] [51]

See also

Notes

  1. ^ Independantly from the alkalinity factor and considering only the saline content, species richness (number of species present, or diversity) in highly saline lakes is surprisingly high, often rivaling that of freshwater ecosystems. Salinity is the primary factor regulating the diversification of bacteria and archaea, including the functional gene lineages involved in the denitrification pathway or the hydrolysis of chitin. [16]
  2. ^ These molecular methods are based on DNA extracted directly from the environment and thus do not require microorganisms to be cultured. This is a major advantage, as culturing of novel microorganisms is a laborious technique known to seriously bias the outcome of diversity studies, since only about one in a hundred organisms can be cultured using standard techniques. [22] For microorganisms, the phylogenetic marker gene small subunit (SSU) ribosomal RNA is typically targeted, due to its particular properties such as existence in all cellular organisms and ability to be used as a "molecular clock" to trace the evolutionary history of an organism. [23]
  3. ^ For Lake Chitu in Ethiopia, see also:
    • Assaye, Hirut; Belay, Amha; Desse, Gulelat Haki; Gray, David (2018). "Seasonal variation in the nutrient profile of Arthrospira fusiformis biomass harvested from an Ethiopian soda lake, Lake Chitu" (PDF). Journal of Applied Phycology. 30 (3): 1597–1606. doi: 10.1007/s10811-017-1359-0. Retrieved 2024-07-14.
    • Ogato, Tadesse; Kifle, Demeke; Fetahi, Tadesse; Sitotaw, Baye (2014). "Evaluation of growth and biomass production of Arthrospira (Spirulina) fusiformis in laboratory cultures using waters from the Ethiopian soda lakes Chitu and Shala". Journal of Applied Phycology. doi: 10.1007/s10811-014-0251-4. Retrieved 2024-07-14.

References

  1. ^ a b c d e f g h i j Grant 2006.
  2. ^ Kempe & Kazmierczak 2011a, p. 825, fig. 1.
  3. ^ a b Kempe & Kazmierczak 2011a, p. 824.
  4. ^ Andreote et al. 2018, abstract.
  5. ^ a b Jônatas, Abrahão; Silva, Lorena; Silva, Ludmila Santos; Khalil, Jacques Yaacoub Bou; Rodrigues, Rodrigo; Arantes, Thalita; Assis, Felipe; Boratto, Paulo; Andrade, Miguel; Kroon, Erna Geessien; Ribeiro, Bergmann; Bergier, Ivan; Seligmann, Herve; Ghigo, Eric; Colson, Philippe; Levasseur, Anthony; Kroemer, Guido; Raoult, Didier; Scola, Bernard La (2018). "Tailed giant Tupanvirus possesses the most complete translational apparatus of the known virosphere". Nature Communications. 9: 749. doi: 10.1038/s41467-018-03168-1. PMC  5829246. PMID  29487281.{{ cite journal}}: CS1 maint: PMC format ( link)
  6. ^ a b c d Kempe & Kazmierczak 2011a, p. 825.
  7. ^ Kempe & Kazmierczak 2011a, p. 825-826.
  8. ^ Kempe & Kazmierczak 2011a, p. 827.
  9. ^ a b Kempe & Kazmierczak 2011a, p. 825, 826, 828.
  10. ^ Kempe & Kazmierczak 2011a, p. 824, 828.
  11. ^ a b Kempe & Kazmierczak 2011a, p. 828.
  12. ^ Melack & Kilham 1974.
  13. ^ a b Byrne et al. 2024.
  14. ^ a b c d e f g h i j Lanzén et al. 2013.
  15. ^ Sorokin, Banciu & Muyzer 2015.
  16. ^ a b Wang J (2011). "Do patterns of bacterial diversity along salinity gradients differ from those observed for macroorganisms?". PLOS ONE. 6 (11): e27597. Bibcode: 2011PLoSO...627597W. doi: 10.1371/journal.pone.0027597. PMC  3220692. PMID  22125616.
  17. ^ a b c Barberán, A.; Casamayor, E. O. (2010). "Euxinic Freshwater Hypolimnia Promote Bacterial Endemicity in Continental Areas". Microbial Ecology. 61 (2): 465–472. doi: 10.1007/s00248-010-9775-6. PMID  21107832. S2CID  6985343.
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