Microbialites
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Microbialite is a benthic sedimentary deposit made of carbonate mud (particle diameter < 5 μm) that is formed with the mediation of microbes. The constituent carbonate mud is a type of
automicrite ''Automicrite'' is autochthonous micrite, that is, a Carbonate minerals, carbonate mud precipitated in situ (no transporting) and made up of fine-grained calcite or aragonite micron-sized crystals. It precipitates on the sea floor or within the sedi ...
, or authigenic
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 ...
mud, and therefore it precipitates in situ instead of being transported and deposited. Being formed in situ, a microbialite can be seen as a type of boundstone where reef builders are microbes, and precipitation of carbonate is biotically induced instead of forming tests, shells or skeletons. Microbialites can also be defined as
microbial mat A microbial mat is a multi-layered sheet of microorganisms, mainly bacteria and archaea, or bacteria alone. Microbial mats grow at interfaces between different types of material, mostly on submerged or moist surfaces, but a few survive in deserts ...
s with lithification capacity.
Bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
can precipitate carbonate both in shallow and in deep water (except for Cyanobacteria) and so microbialites can form regardless of the sunlight. Microbialites are the foundation of many
lacustrine A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much larger ...
ecosystems, such as the biosystem of the Great Salt Lake with its millions of migratory birds or, serving in the Alchichica Lake as nurseries for axolotl (''Ambystoma taylori'') and a variety of fish. Microbialites were very important to the formation of Precambrian and Phanerozoic limestones in many different environments, marine and not. The best age for stromatolites was from 2800 Ma to 1000 Ma where
stromatolites Stromatolites () or stromatoliths () are layered sedimentary formations (microbialite) that are created mainly by photosynthetic microorganisms such as cyanobacteria, sulfate-reducing bacteria, and Pseudomonadota (formerly proteobacteria). Th ...
were the main constituents of
carbonate platform A carbonate platform is a sedimentary body which possesses topographic relief, and is composed of autochthonic calcareous deposits. Platform growth is mediated by sessile organisms whose skeletons build up the reef or by organisms (usually micr ...
s


Classification

Microbialites can have three different fabrics: * Stromatolitic: microbialite layered, laminated or agglutinated to form a
stromatolite Stromatolites () or stromatoliths () are layered sedimentary formations ( microbialite) that are created mainly by photosynthetic microorganisms such as cyanobacteria, sulfate-reducing bacteria, and Pseudomonadota (formerly proteobacteria). T ...
. * Thrombolitic: microbialite with a clotted peloidal fabric if observed with a petrographic microscope. The density of
peloids Peloids are allochems that are composed of micrite, irrespective of size, shape, or origin. The two primary types of peloids are pellets and intraclasts Intraclasts are irregularly shaped grains that form by syndepositional erosion of partially ...
is variable. At the scale of the hand sample, the rock shows a dendritic fabric, and can be named
thrombolite Thrombolites (from Ancient Greek θρόμβος ''thrómbos'' meaning " clot" and λῐ́θος ''líthos'' meaning "stone") are clotted accretionary structures formed in shallow water by the trapping, binding, and cementation of sedimentary g ...
. * Leiolitic: a microbialite with no layering nor clotted peloidal fabric. It is only made of a dense
automicrite ''Automicrite'' is autochthonous micrite, that is, a Carbonate minerals, carbonate mud precipitated in situ (no transporting) and made up of fine-grained calcite or aragonite micron-sized crystals. It precipitates on the sea floor or within the sedi ...
.


Evolution

Microbialites played an important role in the evolution of the Earth's atmosphere, since they were ancestral niches where the first microbial metabolisms capable of releasing oxygen arose. Microbialites saturated coastal systems and later the primitive atmosphere with oxygen, changing it from a reduced state to an oxidized state. The fossil microbialites (also called stromatolites) of the Precambrian and Phanerozoic are one of the first evidences of communal life. The oldest microbialites are dated at 3.5 billion years. Fossil evidence suggests that microbialite-producing organisms were a very abundant life form from the early Archaean to the late Proterozoic, until their communities decreased due to the predation of
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly ...
and other eukaryotic microorganisms.


Formation of microbialites

The formation of microbialites is complex and is a continuous process of
precipitation In meteorology, precipitation is any product of the condensation of atmospheric water vapor that falls under gravitational pull from clouds. The main forms of precipitation include drizzle, rain, sleet, snow, ice pellets, graupel and hail. ...
and
dissolution Dissolution may refer to: Arts and entertainment Books * ''Dissolution'' (''Forgotten Realms'' novel), a 2002 fantasy novel by Richard Lee Byers * ''Dissolution'' (Sansom novel), a 2003 historical novel by C. J. Sansom Music * Dissolution, in mu ...
, where different microbial metabolisms are coupled and a high saturation index (SI) of ions in water is present. Microbialites have two possible genesis mechanisms: 1) Accretion / entrapment: when microorganisms actively trap organic matter, debris or mineral material through extracellular polymeric substances (EPS) . 2) Precipitation: it can be due to inorganic deposition, sedimentation or the passive influence of microbial metabolisms. There can also be precipitation due to saturation of the microenvironment when extracellular polymeric substances are rapidly degraded, causing ion saturation.


Modern microbialites distribution

Living modern microbialites (less than 20,000 years old) are rare and can be found confined to places such as: *
Crater lake Crater Lake ( Klamath: ''Giiwas'') is a volcanic crater lake in south-central Oregon in the western United States. It is the main feature of Crater Lake National Park and is famous for its deep blue color and water clarity. The lake partly fill ...
s: Blue Lake (Australia), Lake Satonda (
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
), Lake Dziani,
Lake Alchichica The Oriental Basin, also known as the Libres-Oriental Basin, Oriental-Serdán Basin or San Juan Plains (in Spanish, ''Llanos de San Juan'' or ''Cuenca de Libres-Oriental)'' is an endorheic basin in east-central Mexico. It covers an area of 4,958.60 ...
(Mexico), Lake Vai Lahi and Lake Vai Sii (
Tonga Tonga (, ; ), officially the Kingdom of Tonga ( to, Puleʻanga Fakatuʻi ʻo Tonga), is a Polynesian country and archipelago. The country has 171 islands – of which 45 are inhabited. Its total surface area is about , scattered over in ...
),
Lake Salda Lake Salda is a mid-size crater lake in southwestern Turkey, within the boundaries of Yeşilova district of Burdur Province. It lies at a distance of about fifty kilometers to the west from the province seat Burdur. Lake Salda is often included ...
(Turkey) * Saline / hypersaline lakes / lagoons: Pyramid Lake and Great Salt Lake (United States),
Lake Van Lake Van ( tr, Van Gölü; hy, Վանա լիճ, translit=Vana lič̣; ku, Gola Wanê) is the largest lake in Turkey. It lies in the far east of Turkey, in the provinces of Van and Bitlis in the Armenian highlands. It is a saline soda lake ...
(Turkey), Brava Lagoon and Tebinquicho Lagoon (
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
),
Lake Van Lake Van ( tr, Van Gölü; hy, Վանա լիճ, translit=Vana lič̣; ku, Gola Wanê) is the largest lake in Turkey. It lies in the far east of Turkey, in the provinces of Van and Bitlis in the Armenian highlands. It is a saline soda lake ...
(Turkey) *
Alkaline lake A soda lake or alkaline lake is a lake on the strongly alkaline side of neutrality, typically with a pH, pH value between 9 and 12. They are characterized by high concentrations of carbonate salts, typically sodium carbonate (and related salt com ...
s: Lake Thetis (Autralia), Lake Sarmiento (Chile), Lake Nuoertu and Lake Huhejaran (China),
Mono Lake Mono Lake ( ) is a saline soda lake in Mono County, California, formed at least 760,000 years ago as a terminal lake in an endorheic basin. The lack of an outlet causes high levels of salts to accumulate in the lake which make its water a ...
(United States), Lake Turkana (Kenya), Lake Petukhovskoe (Russia) * Freshwater lakes /
lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') ...
s: Lagoa Salgada (Brazil), Laguna Negra, Catamarca (Argentina),
Lagunas de Ruidera The Lagunas de Ruidera are a group of small lakes in the Campo de Montiel, Castilla-La Mancha, between Albacete Province, and Ciudad Real Province, Spain. Most of the lakes are interconnected and their total water amount may reach 23.06 hm3 ...
(Spain),
Bacalar Bacalar () is the municipal seat and largest city in Bacalar Municipality (until 2011 a part of Othón P. Blanco Municipality) in the Mexican state of Quintana Roo, about north of Chetumal, at 18° 40' 37" N, 88° 23' 43" W. In the 2010 census ...
(Mexico),
Lake Richmond Lake Richmond (Aboriginal Australian name: Naragebup) is a freshwater lake in Rockingham, Western Australia. It is approximately , with an area of , and is deep in the centre. It is believed to be named after the London borough. It is part of ...
(Australia),
Pavilion Lake Pavilion Lake is a freshwater lake located in Marble Canyon, British Columbia, Canada home to colonies of freshwater microbialites. Location and Local Communities It is located between the towns of Lillooet and Cache Creek (29.44 kilometres WN ...
(Canada), Green Lake (United States) Alkaline pools: Four swamp blue pools (Mexico) Abandoned open mines: Clinton Creek (Canada),
Rio Tinto Rio Tinto, meaning "red river", may refer to: Businesses * Rio Tinto (corporation), an Anglo-Australian multinational mining and resources corporation ** Rio Tinto Alcan, based in Canada ** Rio Tinto Borax in America *** Rio Tinto Borax Mine, ...
(Spain) * Marine / Estuary /
Estuary An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environm ...
Systems: Shark Bay, Australia, Highbourne Cay (
Bahamas The Bahamas (), officially the Commonwealth of The Bahamas, is an island country within the Lucayan Archipelago of the West Indies in the North Atlantic. It takes up 97% of the Lucayan Archipelago's land area and is home to 88% of the ar ...
),
Tikehau Tikehau (meaning ''Peaceful Landing'' in TuamotuanOfficial Tikehau Tourism Site< ...
( French Polynesia), Cayo Coco (Cuba),
Lake Clifton, Western Australia Lake Clifton is a small town located on the east side of the lake of the same name in the Peel region of Western Australia just off the Old Coast Road, between Mandurah and Bunbury at the north end of the Yalgorup National Park. At the 2016 ...
.


Composition

Microbialites are made up of layers made up of an organic component and another mineral. The organic component is an elaborate microbial mat where different communities of microorganisms interact with different metabolisms and create a micro-niche where oxygenic and anoxygenic phototrophic organisms coexist, nitrogen fixers, sulfur reducers, methaneotrophs, methanogens, iron oxidizers, and an infinity of heterotrophic decomposers. The mineral component is composed of carbonates, generally calcium carbonate or magnesium carbonates such as
hydromagnesite Hydromagnesite is a hydrated magnesium carbonate mineral with the formula Mg5(CO3)4(OH)2·4H2O. It generally occurs associated with the weathering products of magnesium containing minerals such as serpentine or brucite. It occurs as incrustations ...
, although there may also be sintered silicones, that is, silicates and include mineral forms of sulfur, iron (pyrite) or phosphorus. Carbonate is usually a type of autogenic automicrite, therefore it precipitates in situ. Microbialites can be viewed as a type of biogenic sedimentary rock where the reef builders are microbes and carbonate precipitation is induced. Microorganisms can precipitate carbonate in both shallow and deep waters


Microbes that produce microbialites

A broad number of studies have analyzed the diversity of microorganisms living at the surface of microbialites. Very often, this diversity is very high and includes bacteria, archaea and eukaryotes. While the phylogenetic diversity of these microbial communities is pretty well assessed using molecular biology, the identity of the organisms contributing to carbonate formation remains uncertain. Interestingly, some microorganisms seem to be present in microbialites forming in several different lakes, defining a core microbiome. Microbes that precipitate carbonate to build microbialites are mostly
prokaryote 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 & Conne ...
s, which include bacteria and archaea. The best known carbonate-producing bacteria are Cyanobacteria and
Sulfate-reducing bacteria Sulfate-reducing microorganisms (SRM) or sulfate-reducing prokaryotes (SRP) are a group composed of sulfate-reducing bacteria (SRB) and sulfate-reducing archaea (SRA), both of which can perform anaerobic respiration utilizing sulfate () as termina ...
. Additional bacteria may play a prominent role, such as bacteria performing anoxygenic photosythes is. Archaea are often
extremophile An extremophile (from Latin ' meaning "extreme" and Greek ' () meaning "love") is an organism that is able to live (or in some cases thrive) in extreme environments, i.e. environments that make survival challenging such as due to extreme temper ...
s and thus live in remote environments where other organisms cannot live, such as white smokers at the bottom of the oceans. Eukaryotic microbes, instead, produce less carbonate than prokaryotes.


Interest in studying microbialites

There is great interest in studying fossil microbialites in the field of paleontology since they provide relevant data on paleoclimate and function as bioclimatic indicators. There is also an interest in studying them in the field of astrobiology, as they are one of the first forms of life, one would expect to find evidence of these structures on other planets. The study of modern microbialites can provide relevant information and serve as environmental indicators for the management and conservation of protected natural areas. Due to their ability to form minerals and precipitate detrital material, biotechnological and bioremediation applications have been suggested in aquatic systems for carbon dioxide sequestration, since microbialites can function as carbon sinks.


References

{{reflist Limestone Sedimentology Stromatolites