Lomagundi Event
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The Great Oxidation Event (GOE), also called the Great Oxygenation Event, the Oxygen Catastrophe, the Oxygen Revolution, the Oxygen Crisis, or the Oxygen Holocaust, was a time interval during the Paleoproterozoic era when the Earth's atmosphere and the shallow ocean first experienced a rise in the amount of oxygen. This began approximately 2.460–2.426 Ga (billion years) ago, during the Siderian period, and ended approximately 2.060 Ga, during the Rhyacian. Geological, isotopic, and chemical evidence suggests that biologically-produced molecular oxygen ( dioxygen, O2) started to accumulate in Earth's atmosphere and changed it from a weakly reducing atmosphere practically free of oxygen into an oxidizing atmosphere containing abundant oxygen, with oxygen levels being as high as 10% of their present atmospheric level by the end of the GOE. The sudden injection of toxic oxygen into an anaerobic biosphere may have caused the extinction of many existing anaerobic species on Earth. Although the event is inferred to have constituted a mass extinction, due in part to the great difficulty in surveying microscopic species' abundances, and in part to the extreme age of fossil remains from that time, the Great Oxidation Event is typically not counted among conventional lists of "great extinctions", which are implicitly limited to the Phanerozoic eon. In any case, isotope geochemical data from sulfate minerals have been interpreted to indicate a decrease in the size of the biosphere of >80% associated with changes in nutrient supplies at the end of the GOE. The GOE is inferred to have been caused by
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
producing the oxygen, which may have enabled the subsequent development of multicellular life-forms.


The early atmosphere

The composition of the Earth's earliest atmosphere is not known with certainty. However, the bulk was likely nitrogen, , and carbon dioxide, , which are also the predominant nitrogen- and carbon-bearing gases produced by volcanism today. These are relatively inert gases. Oxygen, , meanwhile, was present in the atmosphere at just 0.001% of its present atmospheric level. The Sun shone at about 70% of its current brightness 4 billion years ago, but there is strong evidence that liquid water existed on Earth at the time. A warm Earth, in spite of a faint Sun, is known as the faint young Sun paradox. Either carbon dioxide levels were much higher at the time, providing enough of a greenhouse effect to warm the Earth, or other
greenhouse gas A greenhouse gas (GHG or GhG) is a gas that Absorption (electromagnetic radiation), absorbs and Emission (electromagnetic radiation), emits radiant energy within the thermal infrared range, causing the greenhouse effect. The primary greenhouse ...
es were present. The most likely such gas is methane, , which is a powerful greenhouse gas and was produced by early forms of life known as methanogens. Scientists continue to research how the Earth was warmed before life arose. An atmosphere of and with trace amounts of , , carbon monoxide (), and hydrogen (), is described as a weakly reducing atmosphere. Such an atmosphere contains practically no oxygen. The modern atmosphere contains abundant oxygen, making it an oxidizing atmosphere. The rise in oxygen is attributed to photosynthesis by
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
, which are thought to have evolved as early as 3.5 billion years ago. The current scientific understanding of when and how the Earth's atmosphere changed from a weakly reducing to a strongly oxidizing atmosphere largely began with the work of the American geologist Preston Cloud in the 1970s. Cloud observed that detrital sediments older than about 2 billion years ago contained grains of pyrite, uraninite, and siderite, all minerals containing reduced forms of iron or uranium that are not found in younger sediments because they are rapidly oxidized in an oxidizing atmosphere. He further observed that continental red beds, which get their color from the oxidized ( ferric) mineral
hematite Hematite (), also spelled as haematite, is a common iron oxide compound with the formula, Fe2O3 and is widely found in rocks and soils. Hematite crystals belong to the rhombohedral lattice system which is designated the alpha polymorph of . ...
, began to appear in the geological record at about this time. Banded iron formation largely disappears from the geological record at 1.85 billion years ago, after peaking at about 2.5 billion years ago. Banded iron formation can form only when abundant dissolved ferrous iron is transported into depositional basins, and an oxygenated ocean blocks such transport by oxidizing the iron to form insoluble ferric iron compounds. The end of the deposition of banded iron formation at 1.85 billion years ago is therefore interpreted as marking the oxygenation of the deep ocean.
Heinrich Holland Heinrich Dieter 'Dick' Holland (May 27, 1927 – May 21, 2012) was an emeritus professor in the Earth and Planetary Sciences department of Harvard University. He made major contributions to the understanding of the Earth's geochemistry, especia ...
further elaborated these ideas through the 1980s, placing the main time interval of oxygenation between 2.2 and 1.9 billion years ago, and they continue to shape the current scientific understanding. Constraining the onset of atmospheric oxygenation has proven particularly challenging for geologists and geochemists. While there is a widespread consensus that initial oxygenation of the atmosphere happened sometime during the first half of the Paleoproterozoic, there is disagreement on the exact timing of this event. Scientific publications between 2016 to 2022 have differed in the inferred timing of the onset of atmospheric oxygenation by approximately 500 million years, with estimates ranging from as early as 2.7 Ga to as late as 2.225 Ga. This is in large part due to an incomplete
sedimentary record The geologic record in stratigraphy, paleontology and other natural sciences refers to the entirety of the layers of rock strata. That is, deposits laid down by volcanism or by deposition of sediment derived from weathering detritus (clays, sand ...
for the Paleoproterozoic (e.g., because of
subduction Subduction is a geological process in which the oceanic lithosphere is recycled into the Earth's mantle at convergent boundaries. Where the oceanic lithosphere of a tectonic plate converges with the less dense lithosphere of a second plate, the ...
and metamorphism), uncertainties in depositional ages for many ancient sedimentary units, and uncertainties related to the interpretation of different geological/geochemical proxies. While the effects of an incomplete geological record have been discussed and quantified in the field of paleontology for several decades, particularly with respect to the evolution and extinction of organisms ( the Signor-Lipps Effect), this is rarely quantified when considering geochemical records, and may therefore lead to uncertainties for scientists studying the timing of atmospheric oxygenation.


Geological evidence

Evidence for the Great Oxidation Event is provided by a variety of petrological and geochemical markers that define this geological event.


Continental indicators

Paleosols, detrital grains, and
red beds Red beds (or redbeds) are sedimentary rocks, typically consisting of sandstone, siltstone, and shale, that are predominantly red in color due to the presence of ferric oxides. Frequently, these red-colored sedimentary strata locally contain ...
are evidence of low-level oxygen. Paleosols (fossil soils) older than 2.4 billion years old have low iron concentrations that suggest anoxic weathering. Detrital grains composed of pyrite, siderite, and uraninite (redox-sensitive detrital minerals) are found in sediments older than ca. 2.4 Ga. These minerals are only stable under low oxygen conditions, and so their occurrence as detrital minerals in
fluvial In geography and geology, fluvial processes are associated with rivers and streams and the deposits and landforms created by them. When the stream or rivers are associated with glaciers, ice sheets, or ice caps, the term glaciofluvial or fluviog ...
and deltaic sediments are widely interpreted as evidence of an anoxic atmosphere. In contrast to redox-sensitive detrital minerals are
red beds Red beds (or redbeds) are sedimentary rocks, typically consisting of sandstone, siltstone, and shale, that are predominantly red in color due to the presence of ferric oxides. Frequently, these red-colored sedimentary strata locally contain ...
, red-colored sandstones that are coated with hematite. The occurrence of red beds indicates that there was sufficient oxygen to oxidize iron to its ferric state, and represent a marked contrast to sandstones deposited under anoxic conditions which are often beige, white, grey, or green.


Banded iron formation (BIF)

Banded iron formations are composed of thin alternating layers of chert (a fine-grained form of silica) and iron oxides, magnetite and hematite. Extensive deposits of this rock type are found around the world, almost all of which are more than 1.85 billion years old and most of which were deposited around 2.5 billion years ago. The iron in banded iron formation is partially oxidized, with roughly equal amounts of ferrous and ferric iron. Deposition of banded iron formation requires both an anoxic deep ocean capable of transporting iron in soluble ferrous form, and an oxidized shallow ocean where the ferrous iron is oxidized to insoluble ferric iron and precipitates onto the ocean floor. The deposition of banded iron formation before 1.8 billion years ago suggests the ocean was in a persistent ferruginous state, but deposition was episodic and there may have been significant intervals of euxinia.


Iron speciation

Black laminated
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especial ...
s, rich in organic matter, are often regarded as a marker for anoxic conditions. However, the deposition of abundant organic matter is not a sure indication of anoxia, and burrowing organisms that destroy lamination had not yet evolved during the time frame of the Great Oxygenation Event. Thus laminated black shale by itself is a poor indicator of oxygen levels. Scientists must look instead for geochemical evidence of anoxic conditions. These include ferruginous anoxia, in which dissolved ferrous iron is abundant, and euxinia, in which
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The unde ...
is present in the water. Examples of such indicators of anoxic conditions include the degree of pyritization (DOP), which is the ratio of iron present as pyrite to the total reactive iron. Reactive iron, in turn, is defined as iron found in oxides and oxyhydroxides, carbonates, and reduced sulfur minerals such as pyrites, in contrast with iron tightly bound in silicate minerals. A DOP near zero indicates oxidizing conditions, while a DOP near 1 indicates euxinic conditions. Values of 0.3 to 0.5 are transitional, suggesting anoxic bottom mud under an oxygenated ocean. Studies of the Black Sea, which is considered a modern model for ancient anoxic ocean basins, indicate that high DOP, a high ratio of reactive iron to total iron, and a high ratio of total iron to aluminum are all indicators of transport of iron into a euxinic environment. Ferruginous anoxic conditions can be distinguished from euxenic conditions by a DOP less than about 0.7. The currently available evidence suggests that the deep ocean remained anoxic and ferruginous as late as 580 million years ago, well after the Great Oxygenation Event, remaining just short of euxenic during much of this interval of time. Deposition of banded iron formation ceased when conditions of local euxenia on continental platforms and shelves began precipitating iron out of upwelling ferruginous water as pyrite.


Isotopes

Some of the most persuasive evidence for the Great Oxidation Event is provided by the mass-independent fractionation (MIF) of sulfur. The chemical signature of the MIF of sulfur is found prior to 2.4–2.3 billion years ago but disappears thereafter. The presence of this signature all but eliminates the possibility of an oxygenated atmosphere. Different isotopes of a chemical element have slightly different atomic masses. Most of the differences in geochemistry between isotopes of the same element scale with this mass difference. These include small differences in molecular velocities and diffusion rates, which are described as mass-dependent fractionation processes. By contrast, mass-independent fractionation describes processes that are not proportional to the difference in mass between isotopes. The only such process likely to be significant in the geochemistry of sulfur is photodissociation. This is the process in which a molecule containing sulfur is broken up by solar ultraviolet (UV) radiation. The presence of a clear MIF signature for sulfur prior to 2.4 billion years ago shows that UV radiation was penetrating deep into the Earth's atmosphere. This in turn rules out an atmosphere containing more than traces of oxygen, which would have produced an ozone layer that shielded the lower atmosphere from UV radiation. The disappearance of the MIF signature for sulfur indicates the formation of such an ozone shield as oxygen began to accumulate in the atmosphere. Mass-dependent fractionation of sulphur also indicates the presence of oxygen in that oxygen is required to facilitate repeated redox cycling of sulphur. Mass-dependent fractionation also provides clues to the Great Oxygenation Event. For example, oxidation of manganese in surface rocks by atmospheric oxygen leads to further reactions that oxidize chromium. The heavier 53Cr is oxidized preferentially over the lighter 52Cr, and the soluble oxidized chromium carried into the ocean shows this enhancement of the heavier isotope. The chromium isotope ratio in banded iron formation suggests small but significant quantities of oxygen in the atmosphere before the Great Oxidation Event, and a brief return to low oxygen abundance 500 million years after the Great Oxidation Event. However, the chromium data may conflict with the sulfur isotope data, which calls the reliability of the chromium data into question. It is also possible that oxygen was present earlier only in localized "oxygen oases". Since chromium is not easily dissolved, its release from rocks requires the presence of a powerful acid such as
sulfuric acid Sulfuric acid (American spelling and the preferred IUPAC name) or sulphuric acid ( Commonwealth spelling), known in antiquity as oil of vitriol, is a mineral acid composed of the elements sulfur, oxygen and hydrogen, with the molecular formu ...
(H2SO4) which may have formed through bacterial oxidation of pyrite. This could provide some of the earliest evidence of oxygen-breathing life on land surfaces. Other elements whose mass-dependent fractionation may provide clues to the Great Oxidation Event include carbon, nitrogen, transitional metals such as
molybdenum Molybdenum is a chemical element with the symbol Mo and atomic number 42 which is located in period 5 and group 6. The name is from Neo-Latin ''molybdaenum'', which is based on Ancient Greek ', meaning lead, since its ores were confused with lea ...
and iron, and non-metal elements such as selenium.


Fossils and biomarkers (chemical fossils)

While the Great Oxidation Event is generally thought to be a result of oxygenic photosynthesis by ancestral cyanobacteria, the presence of cyanobacteria in the Archaean before the Great Oxidation Event is a highly controversial topic. Structures that are claimed to be fossils of cyanobacteria exist in rock as old as 3.5 billion years old These include microfossils of supposedly cyanobacterial cells and macrofossils called stromatolites, which are interpreted as colonies of microbes, including cyanobacteria, with characteristic layered structures. Modern stromatolites, which can only be seen in harsh environments such as
Shark Bay Shark Bay (Malgana: ''Gathaagudu'', "two waters") is a World Heritage Site in the Gascoyne region of Western Australia. The http://www.environment.gov.au/heritage/places/world/shark-bay area is located approximately north of Perth, on the ...
in Western Australia, are associated with cyanobacteria and thus fossil stromatolites had long been interpreted as the evidence for cyanobacteria. However, it has increasingly been inferred that at least some of these Archaean fossils were generated abiotically or produced by non-cyanobacterial phototrophic bacteria. Additionally, Archaean sedimentary rocks were once found to contain biomarkers, also known as chemical fossils, interpreted as fossilized membrane lipids from cyanobacteria and
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s. For example, traces of 2α-methylhopanes and steranes that are thought to be derived from cyanobacteria and eukaryotes, respectively, were found in the Pilbara of Western Australia.
Sterane Sterane (cyclopentanoperhydrophenanthrenes) compounds are a class of tetracyclic compounds derived from steroids or sterols via diagenetic and catagenetic degradation and saturation. Steranes have an androstane skeleton with a side chain at carb ...
s are diagenetic products of sterols, which are biosynthesized utilizing molecular oxygen. Thus, steranes can additionally serve as an indicator of oxygen in the atmosphere. However, these biomarker samples have since been shown to have been contaminated and so the results are no longer accepted.


Other indicators

Some elements in marine sediments are sensitive to different levels of oxygen in the environment such as the transition metals
molybdenum Molybdenum is a chemical element with the symbol Mo and atomic number 42 which is located in period 5 and group 6. The name is from Neo-Latin ''molybdaenum'', which is based on Ancient Greek ', meaning lead, since its ores were confused with lea ...
and rhenium. Non-metal elements such as selenium and iodine are also indicators of oxygen levels.


Hypotheses

The ability to generate oxygen via photosynthesis likely first appeared in the ancestors of cyanobacteria. These organisms evolved at least 2.45–2.32 billion years ago, and probably as early as 2.7 billion years ago or earlier. However, oxygen remained scarce in the atmosphere until around 2.0 billion years ago, and banded iron formation continued to be deposited until around 1.85 billion years ago. Given the rapid multiplication rate of cyanobacteria under ideal conditions, an explanation is needed for the delay of at least 400 million years between the evolution of oxygen-producing photosynthesis and the appearance of significant oxygen in the atmosphere. Hypotheses to explain this gap must take into consideration the balance between oxygen sources and oxygen sinks. Oxygenic photosynthesis produces organic carbon that must be segregated from oxygen to allow oxygen accumulation in the surface environment, otherwise the oxygen back-reacts with the organic carbon and does not accumulate. The burial of organic carbon, sulfide, and minerals containing ferrous iron (Fe) is a primary factor in oxygen accumulation. When organic carbon is buried without being oxidized, the oxygen is left in the atmosphere. In total, the burial of organic carbon and pyrite today creates of O per year. This creates a net O flux from the global oxygen sources. The rate of change of oxygen can be calculated from the difference between global sources and sinks. The oxygen sinks include reduced gases and minerals from volcanoes, metamorphism and weathering. The GOE started after these oxygen-sink fluxes and reduced-gas fluxes were exceeded by the flux of O2 associated with the burial of reductants, such as organic carbon. For the weathering mechanisms, of O per year today goes to the sinks composed of reduced minerals and gases from volcanoes, metamorphism, percolating seawater and heat vents from the seafloor. On the other hand, of O per year today oxidizes reduced gases in the atmosphere through photochemical reaction. On the early Earth, there was visibly very little oxidative weathering of continents (e.g., a lack of
red beds Red beds (or redbeds) are sedimentary rocks, typically consisting of sandstone, siltstone, and shale, that are predominantly red in color due to the presence of ferric oxides. Frequently, these red-colored sedimentary strata locally contain ...
) and so the weathering sink on oxygen would have been negligible compared to that from reduced gases and dissolved iron in oceans. Dissolved iron in oceans exemplifies O2 sinks. Free oxygen produced during this time was chemically captured by dissolved iron, converting iron Fe and Fe2+ to magnetite () that is insoluble in water, and sank to the bottom of the shallow seas to create banded iron formations such as the ones found in Minnesota and the Pilbara of Western Australia. It took 50 million years or longer to deplete the oxygen sinks. The rate of photosynthesis and associated rate of organic burial also affect the rate of oxygen accumulation. When land plants spread over the continents in the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
, more organic carbon was buried and likely allowed higher O2 levels to occur. Today, the average time that an O molecule spends in the air before it is consumed by geological sinks is about 2 million years. That residence time is relatively short in geologic time - so in the
Phanerozoic The Phanerozoic Eon is the current geologic eon in the geologic time scale, and the one during which abundant animal and plant life has existed. It covers 538.8 million years to the present, and it began with the Cambrian Period, when anima ...
there must have been feedback processes that kept the atmospheric O level within bounds suitable for animal life.


Evolution by stages

Preston Cloud originally proposed that the first cyanobacteria had evolved the capacity to carry out oxygen-producing photosynthesis, but had not yet evolved enzymes (such as
superoxide dismutase Superoxide dismutase (SOD, ) is an enzyme that alternately catalyzes the dismutation (or partitioning) of the superoxide () radical into ordinary molecular oxygen (O2) and hydrogen peroxide (). Superoxide is produced as a by-product of oxygen me ...
) for living in an oxygenated environment. These cyanobacteria would have been protected from their own poisonous oxygen waste through its rapid removal via the high levels of reduced ferrous iron, Fe(II), in the early ocean. Cloud suggested that the oxygen released by photosynthesis oxidized the Fe(II) to ferric iron, Fe(III), which precipitated out of the
sea water Seawater, or salt water, is water from a sea or ocean. On average, seawater in the world's oceans has a salinity of about 3.5% (35 g/L, 35 ppt, 600 mM). This means that every kilogram (roughly one liter by volume) of seawater has approx ...
to form banded iron formation. Cloud interpreted the great peak in deposition of banded iron formation at the end of the Archean as the signature for the evolution of mechanisms for living with oxygen. This ended self-poisoning and produced a population explosion in the cyanobacteria that rapidly oxygenated the ocean and ended banded iron formation deposition. However, improved dating of Precambrian strata showed that the late Archean peak of deposition was spread out over tens of millions of years, rather than taking place in a very short interval of time following the evolution of oxygen-coping mechanisms. This made Cloud's hypothesis untenable. Most modern interpretations describe the GOE as a long, protracted process that took place over hundreds of millions of years rather than a single abrupt event, with the quantity of atmospheric oxygen yo-yoing in relation to the capacity of oxygen sinks and the productivity of oxygenic photosynthesisers over the course of the GOE. More recently, families of bacteria have been discovered that show no indication of ever having had photosynthetic capability, but which otherwise closely resemble cyanobacteria. These may be descended from the earliest ancestors of cyanobacteria, which only later acquired photosynthetic ability by
lateral gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). H ...
. Based on molecular clock data, the evolution of oxygen-producing photosynthesis may have occurred much later than previously thought, at around 2.5 billion years ago. This reduces the gap between the evolution of oxygen photosynthesis and the appearance of significant atmospheric oxygen.


Nutrient famines

A second possibility is that early cyanobacteria were starved for vital nutrients, and this checked their growth. However, a lack of the scarcest nutrients, iron, nitrogen, and phosphorus, could have slowed, but not prevented, a cyanobacteria population explosion and rapid oxygenation. The explanation for the delay in the oxygenation of the atmosphere following the evolution of oxygen-producing photosynthesis likely lies in the presence of various oxygen sinks on the young Earth.


Nickel famine

Early chemosynthetic organisms likely produced methane, an important trap for molecular oxygen, since methane readily
oxidizes Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a d ...
to carbon dioxide (CO2) and water in the presence of UV radiation. Modern methanogens require nickel as an enzyme cofactor. As the Earth's crust cooled and the supply of volcanic nickel dwindled, oxygen-producing algae began to out-perform methane producers, and the oxygen percentage of the atmosphere steadily increased. From 2.7 to 2.4 billion years ago, the rate of deposition of nickel declined steadily from a level 400 times today's. This nickel famine was somewhat buffered by an uptick in sulphide weathering at the start of the GOE that brought some nickel to the oceans, without which methanogenic organisms would have declined in abundance more precipitously, plunging Earth into even more severe and long-lasting
icehouse Icehouse or ice house may refer to: * Ice house (building), a building where ice is stored * Ice shanty, a shelter for ice fishing also known as an ''Icehouse'' * Ice skating rink, a facility for ice skating. * Ice hockey arena, an area where ice ...
conditions than those seen during the Huronian glaciation.


Increasing flux

Some people suggest that GOE is caused by the increase of the source of oxygen. One hypothesis argues that GOE was the immediate result of photosynthesis, although the majority of scientists suggest that a long-term increase of oxygen is more likely. Several model results show possibilities of long-term increase of carbon burial, but the conclusions are indecisive.


Decreasing sink

In contrast to the increasing flux hypothesis, there are also several hypotheses that attempt to use decrease of sinks to explain the GOE. One theory suggests that the composition of the volatiles from volcanic gases was more oxidized. Another theory suggests that the decrease of metamorphic gases and serpentinization is the main key of GOE. Hydrogen and methane released from metamorphic processes are also lost from Earth's atmosphere over time and leave the crust oxidized. Scientists realized that hydrogen would escape into space through a process called methane photolysis, in which methane decomposes under the action of ultraviolet light in the upper atmosphere and releases its hydrogen. The escape of hydrogen from the Earth into space must have oxidized the Earth because the process of hydrogen loss is chemical oxidation. This process of hydrogen escape required the generation of methane by methanogens, so that methanogens actually helped create the conditions necessary for the oxidation of the atmosphere.


Tectonic trigger

One hypothesis suggests that the oxygen increase had to await tectonically driven changes in the Earth, including the appearance of shelf seas, where reduced organic carbon could reach the sediments and be buried. The newly produced oxygen was first consumed in various chemical reactions in the oceans, primarily with iron. Evidence is found in older rocks that contain massive banded iron formations apparently laid down as this iron and oxygen first combined; most present-day
iron ore Iron ores are rocks and minerals from which metallic iron can be economically extracted. The ores are usually rich in iron oxides and vary in color from dark grey, bright yellow, or deep purple to rusty red. The iron is usually found in the fo ...
lies in these deposits. It was assumed oxygen released from cyanobacteria resulted in the chemical reactions that created rust, but it appears the iron formations were caused by anoxygenic phototrophic iron-oxidizing bacteria, which does not require oxygen. Evidence suggests oxygen levels spiked each time smaller land masses collided to form a super-continent. Tectonic pressure thrust up mountain chains, which eroded releasing nutrients into the ocean that fed photosynthetic cyanobacteria.


Bistability

Another hypothesis posits a model of the atmosphere that exhibits
bistability In a dynamical system, bistability means the system has two stable equilibrium states. Something that is bistable can be resting in either of two states. An example of a mechanical device which is bistable is a light switch. The switch lever ...
: two steady states of oxygen concentration. The state of stable low oxygen concentration (0.02%) experiences a high rate of methane oxidation. If some event raises oxygen levels beyond a moderate threshold, the formation of an
ozone layer The ozone layer or ozone shield is a region of Earth's stratosphere that absorbs most of the Sun's ultraviolet radiation. It contains a high concentration of ozone (O3) in relation to other parts of the atmosphere, although still small in rela ...
shields UV rays and decreases methane oxidation, raising oxygen further to a stable state of 21% or more. The Great Oxygenation Event can then be understood as a transition from the lower to the upper steady states.


Increasing photoperiod

Cyanobacteria tend to consume nearly as much oxygen at night as they produce during the day. However, experiments demonstrate that cyanobacterial mats produce a greater excess of oxygen with longer photoperiods. The rotational period of the Earth was only about six hours shortly after its formation, 4.5 billion years ago, but increased to 21 hours by 2.4 billion years ago, in the Paleoproterozoic. The rotational period increased again, starting 700 million years ago, to its present value of 24 hours. It is possible that each increase in rotational period increased the net oxygen production by cyanobacterial mats, which in turn increased the atmospheric abundance of oxygen.


Consequences of oxygenation

Eventually, oxygen started to accumulate in the atmosphere, with two major consequences. * Oxygen likely oxidized atmospheric methane (a strong
greenhouse gas A greenhouse gas (GHG or GhG) is a gas that Absorption (electromagnetic radiation), absorbs and Emission (electromagnetic radiation), emits radiant energy within the thermal infrared range, causing the greenhouse effect. The primary greenhouse ...
) to carbon dioxide (a weaker one) and water. This weakened the greenhouse effect of the Earth's atmosphere, causing planetary cooling, which has been proposed to have triggered a series of ice ages known as the Huronian glaciation, bracketing an age range of 2.45–2.22 billion years ago. * The increased oxygen concentrations provided a new opportunity for biological diversification, as well as tremendous changes in the nature of chemical interactions between
rocks In geology, rock (or stone) is any naturally occurring solid mass or aggregate of minerals or mineraloid matter. It is categorized by the minerals included, its chemical composition, and the way in which it is formed. Rocks form the Earth's ...
, sand, clay, and other geological substrates and the Earth's air, oceans, and other surface waters. Despite the natural recycling 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 ...
, life had remained energetically limited until the widespread availability of oxygen. The availability of oxygen greatly increased the free energy available to living organisms, with global environmental impacts. For example,
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
evolved after the GOE, giving organisms the energy to exploit new, more complex morphologies interacting in increasingly complex ecosystems, although these did not appear until the late Proterozoic and Cambrian.


Role in mineral diversification

The Great Oxygenation Event triggered an explosive growth in the diversity of minerals, with many elements occurring in one or more oxidized forms near the Earth's surface. It is estimated that the GOE was directly responsible for more than 2,500 of the total of about 4,500 minerals found on Earth today. Most of these new minerals were formed as hydrated and oxidized forms due to dynamic
mantle A mantle is a piece of clothing, a type of cloak. Several other meanings are derived from that. Mantle may refer to: *Mantle (clothing), a cloak-like garment worn mainly by women as fashionable outerwear **Mantle (vesture), an Eastern Orthodox ve ...
and crust processes.


Role in cyanobacteria evolution

In field studies done in Lake Fryxell, Antarctica, scientists found that mats of oxygen-producing cyanobacteria produced a thin layer, one to two millimeters thick, of oxygenated water in an otherwise anoxic environment, even under thick ice. By inference, these organisms could have adapted to oxygen even before oxygen accumulated in the atmosphere. * The evolution of such oxygen-dependent organisms eventually established an equilibrium in the availability of oxygen, which became a major constituent of the atmosphere.


Origin of eukaryotes

It has been proposed that a local rise in oxygen levels due to
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
l photosynthesis in ancient microenvironments was highly toxic to the surrounding biota, and that this selective pressure drove the evolutionary transformation of an
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
l lineage into the first
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s. Oxidative stress involving production of reactive oxygen species (ROS) might have acted in synergy with other environmental stresses (such as ultraviolet radiation and/or
desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...
) to drive selection in an early archaeal lineage towards eukaryosis. This archaeal ancestor may already have had DNA repair mechanisms based on DNA pairing and recombination and possibly some kind of cell fusion mechanism. The detrimental effects of internal ROS (produced by endosymbiont proto-
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
) on the archaeal genome could have promoted the evolution of meiotic sex from these humble beginnings. Selective pressure for efficient DNA repair of oxidative DNA damage may have driven the evolution of eukaryotic sex involving such features as cell-cell fusions, cytoskeleton-mediated chromosome movements and emergence of the nuclear membrane. Thus the evolution of eukaryotic sex and eukaryogenesis were likely inseparable processes that evolved in large part to facilitate DNA repair.


Lomagundi-Jatuli event

The rise in oxygen content was not linear: instead, there was a rise in oxygen content around 2.3 Ga ago, followed by a drop around 2.1 Ga ago. The positive excursion, or more precisely, the carbon isotopic excursion evidencing it, is called the Lomagundi-Jatuli event (LJE) or Lomagundi event, (named for a district of Southern Rhodesia) and the time period has been termed Jatulian; it is believed to be part of the Rhyacian period. In the Lomagundi-Jatuli event, oxygen content reached as high as modern levels, followed by a fall to very low levels during the following stage where black shales were deposited. The negative excursion is called the . Evidence for the Lomagundi-Jatuli event has been found globally: in
Fennoscandia __NOTOC__ Fennoscandia (Finnish language, Finnish, Swedish language, Swedish and no, Fennoskandia, nocat=1; russian: Фенноскандия, Fennoskandiya) or the Fennoscandian Peninsula is the geographical peninsula in Europe, which includes ...
and northern Russia, Scotland, Ukraine,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, the Wyoming Craton in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, Brazil, Uruguay, Gabon, Zimbabwe, South Africa, India, and
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
. Oceans seem to have been oxygenated for some time even after the termination of the isotope excursion itself. It has been hypothesized that
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s first evolved during the LJE. The Lomagundi-Jatuli event coincides with the appearance and subsequent disappearance of curious fossils found in Gabon that have been termed the Francevillian biota, which seem to have been multicellular. This appears to represent a "false start" of multicellular life. The organisms apparently went extinct when the LJE ended, because they are absent in the layers of shale deposited after the LJE.


See also

* Boring Billion – Earth history between 1.8 and 0.8 billion years ago, characterized by tectonic stability, climatic stasis, and a slow biological evolution with very low oxygen levels and no evidence of glaciation * Geological history of oxygen – Timeline of the development of free oxygen in the Earth's oceans and atmosphere * * *
Rare Earth hypothesis In planetary astronomy and astrobiology, the Rare Earth hypothesis argues that the origin of life and the evolution of biological complexity such as sexually reproducing, multicellular organisms on Earth (and, subsequently, human intelligence) ...
– Hypothesis that complex extraterrestrial life is an extremely rare phenomenon * *


References


External links

* {{Extinction Paleoproterozoic Origin of life Oxygen Events in the geological history of Earth Evolution of the biosphere Extinction events Mass extinction timelines Meteorological hypotheses