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Earth's history with time-spans of the eons to scale

The history of Earth concerns the development of planet Earth from its formation to the present day.[1][2] Nearly all branches of natural science have contributed to understanding of the main events of Earth's past, characterized by constant geological change and biological evolution.

The geological time scale (GTS), as defined by international convention,[3] depicts the large spans of time from the beginning of the Earth to the present, and its divisions chronicle some definitive events of Earth history. (In the graphic: Ga means "billion years ago"; Ma, "million years ago".) Earth formed around 4.54 billion years ago, approximately one-third the age of the universe, by accretion from the solar nebula.[4][5][6] Volcanic outgassing probably created the primordial atmosphere and then the ocean, but the early atmosphere contained almost no oxygen. Much of the Earth was molten because of frequent collisions with other bodies which led to extreme volcanism. While the Earth was in its earliest stage (Early Earth), a giant impact collision with a planet-sized body named Theia is thought to have formed the Moon. Over time, the Earth cooled, causing the formation of a solid crust, and allowing liquid water on the surface.

The Hadean eon represents the time before a reliable (fossil) record of life; it began with the formation of the planet and ended 4.0 billion years ago. The following Archean and Proterozoic eons produced the beginnings of life on Earth and its earliest evolution. The succeeding eon is the Phanerozoic, divided into three eras: the Palaeozoic, an era of arthropods, fishes, and the first life on land; the Mesozoic, which spanned the rise, reign, and climactic extinction of the non-avian dinosaurs; and the Cenozoic, which saw the rise of mammals. Recognizable humans emerged at most 2 million years ago, a vanishingly small period on the geological scale.

The earliest undisputed evidence of life on Earth dates at least from 3.5 billion years ago,[7][8][9] during the Eoarchean Era, after a geological crust started to solidify following the earlier molten Hadean Eon. There are microbial mat fossils such as stromatolites found in 3.48 billion-year-old sandstone discovered in Western Australia.[10][11][12] Other early physical evidence of a biogenic substance is graphite in 3.7 billion-year-old metasedimentary rocks discovered in southwestern Greenland[13] as well as "remains of biotic life" found in 4.1 billion-year-old rocks in Western Australia.[14][15] According to one of the researchers, "If life arose relatively quickly on Earth … then it could be common in the universe."[14]

Photosynthetic organisms appeared between 3.2 and 2.4 billion years ago and began enriching the atmosphere with oxygen. Life remained mostly small and microscopic until about 580 million years ago, when complex planet Earth from its formation to the present day.[1][2] Nearly all branches of natural science have contributed to understanding of the main events of Earth's past, characterized by constant geological change and biological evolution.

The geological time scale (GTS), as defined by international convention,[3] depicts the large spans of time from the beginning of the Earth to the present, and its divisions chronicle some definitive events of Earth history. (In the graphic: Ga means "billion years ago"; Ma, "million years ago".) Earth formed around 4.54 billion years ago, approximately one-third the age of the universe, by accretion from the geological time scale (GTS), as defined by international convention,[3] depicts the large spans of time from the beginning of the Earth to the present, and its divisions chronicle some definitive events of Earth history. (In the graphic: Ga means "billion years ago"; Ma, "million years ago".) Earth formed around 4.54 billion years ago, approximately one-third the age of the universe, by accretion from the solar nebula.[4][5][6] Volcanic outgassing probably created the primordial atmosphere and then the ocean, but the early atmosphere contained almost no oxygen. Much of the Earth was molten because of frequent collisions with other bodies which led to extreme volcanism. While the Earth was in its earliest stage (Early Earth), a giant impact collision with a planet-sized body named Theia is thought to have formed the Moon. Over time, the Earth cooled, causing the formation of a solid crust, and allowing liquid water on the surface.

The Hadean eon represents the time before a reliable (fossil) record of life; it began with the formation of the planet and ended 4.0 billion years ago. The following Archean and Proterozoic eons produced the beginnings of life on Earth and its earliest evolution. The succeeding eon is the Phanerozoic, divided into three eras: the Palaeozoic, an era of arthropods, fishes, and the first life on land; the Mesozoic, which spanned the rise, reign, and climactic extinction of the non-avian dinosaurs; and the Cenozoic, which saw the rise of mammals. Recognizable humans emerged at most 2 million years ago, a vanishingly small period on the geological scale.

The earliest undisputed evidence of life on Earth dates at least from 3.5 billion years ago,[7][8][9] during the Eoarchean Era, after a geological crust started to solidify following the earlier molten Hadean Eon. There are microbial mat fossils such as stromatolites found in 3.48 billion-year-old sandstone discovered in Western Australia.[10][11][12] Other early physical evidence of a biogenic substance is graphite in 3.7 billion-year-old metasedimentary rocks discovered in southwestern Greenland[13] as well as "remains of biotic life" found in 4.1 billion-year-old rocks in Western Australia.[14][15] According to one of the researchers, "If life arose relatively quickly on Earth … then it could be common in the universe."[14]

Photosynthetic organisms appeared between 3.2 and 2.4 billion years ago and began enriching the atmosphere with oxygen. Life remained mostly small and microscopic until about 580 million years ago, when complex multicellular life arose, developed over time, and culminated in the Cambrian Explosion about 541 million years ago. This sudden diversification of life forms produced most of the major phyla known today, and divided the Proterozoic Eon from the Cambrian Period of the Paleozoic Era. It is estimated that 99 percent of all species that ever lived on Earth, over five billion,[16] have gone extinct.[17][18] Estimates on the number of Earth's current species range from 10 million to 14 million,[19] of which about 1.2 million are documented, but over 86 percent have not been described.[20] However, it was recently claimed that 1 trillion species currently live on Earth, with only one-thousandth of one percent described.[21]

The Earth's crust has constantly changed since its formation, as has life since its first appearance. Species continue to evolve, taking on new forms, splitting into daughter species, or going extinct in the face of ever-changing physical environments. The process of plate tectonics continues to shape the Earth's continents and oceans and the life they harbor. Human activity is now a dominant force affecting global change, harming the biosphere, the Earth's surface, hydrosphere, and atmosphere with the loss of wild lands, over-exploitation of the oceans, production of greenhouse gases, degradation of the ozone layer, and general degradation of soil, air, and water quality.

In geochronology, time is generally measured in mya (million years ago), each unit representing the period of approximately 1,000,000 years in the past. The history of Earth is divided into four great eons, starting 4,540 mya with the formation of the planet. Each eon saw the most significant changes in Earth's composition, climate and life. Each eon is subsequently divided into eras, which in turn are divided into periods, which are further divided into epochs.

Geologic time scale

The history of the Earth can be organized chronologically according to the geologic time scale, which is split into intervals based on stratigraphic analysis.[2][22] The following four timelines show the geologic time scale. The first shows the entire time from the formation of the Earth to the present, but this gives little space for the most recent eon. Therefore, the second timeline shows an expanded view of the most recent eon. In a similar way, the most recent era is expanded in the third timeline, and the most recent period is expanded in the fourth timeline.

SiderianRhyacianOrosirianStatherianCalymmianEctasianStenianTonianCryogenianEdiacaranEoarcheanPaleoarcheanMesoarcheanThe history of the Earth can be organized chronologically according to the geologic time scale, which is split into intervals based on stratigraphic analysis.[2][22] The following four timelines show the geologic time scale. The first shows the entire time from the formation of the Earth to the present, but this gives little space for the most recent eon. Therefore, the second timeline shows an expanded view of the most recent eon. In a similar way, the most recent era is expanded in the third timeline, and the most recent period is expanded in the fourth timeline.

SiderianRhyacianOrosirianStatherianCalymmianEctasianStenianTonianCryogenianEdiacaranEoarcheanPaleoarcheanMesoarcheanNeoarcheanPaleoproterozoicMesoproterozoicNeoproterozoicPaleozoicMesozoicCenozoicHadeanArcheanProterozoicPhanerozoicPrecambrian
CambrianOrdovicianSilurianDevonianCarboniferousPermianTriassicJurassicCretaceousPaleogeneThe standard model for the formation of the Solar System (including the Earth) is the solar nebula hypothesis.[23] In this model, the Solar System formed from a large, rotating cloud of interstellar dust and gas called the solar nebula. It was composed of hydrogen and helium created shortly after the Big Bang 13.8 Ga (billion years ago) and heavier elements ejected by supernovae. About 4.5 Ga, the nebula began a contraction that may have been triggered by the shock wave from a nearby supernova.[24] A shock wave would have also made the nebula rotate. As the cloud began to accelerate, its angular momentum, gravity, and inertia flattened it into a protoplanetary disk perpendicular to its axis of rotation. Small perturbations due to collisions and the angular momentum of other large debris created the means by which kilometer-sized protoplanets began to form, orbiting the nebular center.[25]

The center of the nebula, not having much angular momentum, collapsed rapidly, the compression heating it until nuclear fusion of hydrogen into helium began. After more contraction, a T Tauri star ignited and evolved into the Sun. Meanwhile, in the outer part of the nebula gravity caused matter to condense around density perturbations and dust particles, and the rest of the protoplanetary disk began separating into rings. In a process known as runaway accretion, successively larger fragments of dust and debris clumped together to form planets.[25] Earth formed in this manner about 4.54 billion years ago (with an uncertainty of 1%)[26][27][4][28] and was largely completed within 10–20 million years.[29] The solar wind of the newly formed T Tauri star cleared out most of the material in the disk that had not already condensed into larger bodies. The same process is expected to produce accretion disks around virtually all newly forming stars in the universe, some of which yield planets.[30]

The proto-Earth grew by accretion until its interior was hot enough to melt the heavy, siderophile metals. Having higher densities than the silicates, these metals sank. This so-called iron catastrophe resulted in the separation of a primitive mantle and a (metallic) core only 10 million years after the Earth began to form, producing the layered structure of Earth and setting up the formation of Earth's magnetic field.[31] J.A. Jacobs [32] was the first to suggest that Earth's inner core—a solid center distinct from the liquid outer core—is freezing and growing out of the liquid outer core due to the gradual cooling of Earth's interior (about 100 degrees Celsius per billion years[33]).

Hadean and Archean Eons

Artist's conception of Hadean Eon Earth, when it was much hotter and inhospitable to all forms of life.

The first eon in Earth's history, the Hadean, begins with the Earth's formation and is followed by the Archean eon at 3.8 Ga.[2]:145 The oldest rocks found on Earth date to about 4.0 Ga, and the oldest detrital zircon crystals in rocks to about 4.4 Ga,[34][35][36] soon after the formation of the Earth's crust and the Earth itself. The giant impact hypothesis for the Moon's formation states that shortly after formation of an initial crust, the proto-Earth was impacted by a smaller protoplanet, which ejected part of the mantle and crust into space and created the Moon.[37][38][39]

From crater counts on other celestial bodies, it is inferred that a period of intense meteorite impacts, called the Late Heavy Bombardment, began about 4.1 Ga, and concluded around 3.8 Ga, at the end of the Hadean.[40] In addition, volcanism was severe due to the large heat flow and geothermal gradient.[41] Nevertheless, detrital zircon crystals dated to 4.4 Ga show evidence of having undergone contact with liquid water, suggesting that the Earth already had oceans or seas at that time.[34]

By the beginning of the Archean, the Earth had cooled significantly. Present life forms could not have survived at Earth's surface, because the Archean atmosphere lacked oxygen hence had no ozone layer to block ultraviolet light. Nevertheless, it is believed t

The center of the nebula, not having much angular momentum, collapsed rapidly, the compression heating it until nuclear fusion of hydrogen into helium began. After more contraction, a T Tauri star ignited and evolved into the Sun. Meanwhile, in the outer part of the nebula gravity caused matter to condense around density perturbations and dust particles, and the rest of the protoplanetary disk began separating into rings. In a process known as runaway accretion, successively larger fragments of dust and debris clumped together to form planets.[25] Earth formed in this manner about 4.54 billion years ago (with an uncertainty of 1%)[26][27][4][28] and was largely completed within 10–20 million years.[29] The solar wind of the newly formed T Tauri star cleared out most of the material in the disk that had not already condensed into larger bodies. The same process is expected to produce accretion disks around virtually all newly forming stars in the universe, some of which yield planets.[30]

The proto-Earth grew by accretion until its interior was hot enough to melt the heavy, siderophile metals. Having higher densities than the silicates, these metals sank. This so-called iron catastrophe resulted in the separation of a primitive mantle and a (metallic) core only 10 million years after the Earth began to form, producing the layered structure of Earth and setting up the formation of Earth's magnetic field.[31] J.A. Jacobs [32] was the first to suggest that Earth's inner core—a solid center distinct from the liquid outer core—is freezing and growing out of the liquid outer core due to the gradual cooling of Earth's interior (about 100 degrees Celsius per billion years[33]).

The first eon in Earth's history, the Hadean, begins with the Earth's formation and is followed by the Archean eon at 3.8 Ga.[2]:145 The oldest rocks found on Earth date to about 4.0 Ga, and the oldest detrital zircon crystals in rocks to about 4.4 Ga,[34][35][36] soon after the formation of the Earth's crust and the Earth itself. The giant impact hypothesis for the Moon's formation states that shortly after formation of an initial crust, the proto-Earth was impacted by a smaller protoplanet, which ejected part of the mantle and crust into space and created the Moon.[37][38][39]

From crater counts on other celestial bodies, it is inferred that a period of intense meteorite impacts, called the Late Heavy Bombardment, began about 4.1 Ga, and concluded around 3.8 Ga, at the end of the Hadean.[40] In addition, volcanism was severe due to the large heat flow and geothermal gradient.[41] Nevertheless, detrital zircon crystals dated to 4.4 Ga show evidence of having undergone contact with liquid water, suggesting that the Earth already had oceans or seas at that time.[34]

By the beginning of the Archean, the Ear

From crater counts on other celestial bodies, it is inferred that a period of intense meteorite impacts, called the Late Heavy Bombardment, began about 4.1 Ga, and concluded around 3.8 Ga, at the end of the Hadean.[40] In addition, volcanism was severe due to the large heat flow and geothermal gradient.[41] Nevertheless, detrital zircon crystals dated to 4.4 Ga show evidence of having undergone contact with liquid water, suggesting that the Earth already had oceans or seas at that time.[34]

By the beginning of the Archean, the Earth had cooled significantly. Present life forms could not have survived at Earth's surface, because the Archean atmosphere lacked oxygen hence had no ozone layer to block ultraviolet light. Nevertheless, it is believed that primordial life began to evolve by the early Archean, with candidate fossils dated to around 3.5 Ga.[42] Some scientists even speculate that life could have begun during the early Hadean, as far back as 4.4 Ga, surviving the possible Late Heavy Bombardment period in hydrothermal vents below the Earth's surface.[43]

Earth's only natural satellite, the Moon, is larger relative to its planet than any other satellite in the Solar System.[nb 1] During the Apollo program, rocks from the Moon's surface were brought to Earth. Radiometric dating of these rocks shows that the Moon is 4.53 ± 0.01 billion years old,[46] formed at least 30 million years after the Solar System.[47] New evidence suggests the Moon formed even later, 4.48 ± 0.02 Ga, or 70–110 million years after the start of the Solar System.[48]

Theories for the formation of the Moon must explain its late formation as well as the following facts. First, the Moon has a low density (3.3 times that of water, compared to 5.5 for the Earth[49]) and a small metallic core. Second, there is virtually no water or other volatiles on the Moon. Third, the Earth and Moon have the same oxygen isotopic signature (relative abundance of the oxygen isotopes). Of the theories proposed to account for these phenomena, one is widely accepted: The giant impact hypothesis proposes that the Moon originated after a body the size of Mars (sometimes named Theia[47]) struck the proto-Earth a glancing blow.[1]:256[50][51]

The collision released about 100 million

Theories for the formation of the Moon must explain its late formation as well as the following facts. First, the Moon has a low density (3.3 times that of water, compared to 5.5 for the Earth[49]) and a small metallic core. Second, there is virtually no water or other volatiles on the Moon. Third, the Earth and Moon have the same oxygen isotopic signature (relative abundance of the oxygen isotopes). Of the theories proposed to account for these phenomena, one is widely accepted: The giant impact hypothesis proposes that the Moon originated after a body the size of Mars (sometimes named Theia[47]) struck the proto-Earth a glancing blow.[1]:256[50][51]

The collision released about 100 million times more energy than the more recent Chicxulub impact that is believed to have caused the extinction of the non-avian dinosaurs. It was enough to vaporize some of the Earth's outer layers and melt both bodies.[50][1]:256 A portion of the mantle material was ejected into orbit around the Earth. The giant impact hypothesis predicts that the Moon was depleted of metallic material,[52] explaining its abnormal composition.[53] The ejecta in orbit around the Earth could have condensed into a single body within a couple of weeks. Under the influence of its own gravity, the ejected material became a more spherical body: the Moon.[54]

Mantle convection, the process that drives plate tectonics, is a result of heat flow from the Earth's interior to the Earth's surface.[55]:2 It involves the creation of rigid tectonic plates at mid-oceanic ridges. These plates are destroyed by subduction into the mantle at subduction zones. During the early Archean (about 3.0 Ga) the mantle was much hotter than today, probably around 1,600 °C (2,910 °F),[56]:82 so convection in the mantle was faster. Although a process similar to present-day plate tectonics did occur, this would have gone faster too. It is likely that during the Hadean and Archean, subduction zones were more common, and therefore tectonic plates were smaller.[1]:258[57]

The initial crust, formed when the Earth's surface first solidified, totally disappeared from a combination of this fast Hadean plate tectonics and the intense impacts of the Late Heavy Bombardment. However, it is thought that it was basaltic in composition, like today's oceanic crust, because little crustal differentiation had yet taken place.[1]:258 The first larger pieces of continental crust, which is a product of differentiation of lighter elements during partial melting in the lower crust, appeared at the end of the Hadean, about 4.0 Ga. What is left of these first small continents are called cratons. These pieces of late Hadean and early Archean crust form the cores around which today's continents grew.The initial crust, formed when the Earth's surface first solidified, totally disappeared from a combination of this fast Hadean plate tectonics and the intense impacts of the Late Heavy Bombardment. However, it is thought that it was basaltic in composition, like today's oceanic crust, because little crustal differentiation had yet taken place.[1]:258 The first larger pieces of continental crust, which is a product of differentiation of lighter elements during partial melting in the lower crust, appeared at the end of the Hadean, about 4.0 Ga. What is left of these first small continents are called cratons. These pieces of late Hadean and early Archean crust form the cores around which today's continents grew.[58]

The oldest rocks on Earth are found in the North American craton of Canada. They are tonalites from about 4.0 Ga. They show traces of metamorphism by high temperature, but also sedimentary grains that have been rounded by erosion during transport by water, showing that rivers and seas existed then.[59] Cratons consist primarily of two alternating types of terranes. The first are so-called greenstone belts, consisting of low-grade metamorphosed sedimentary rocks. These "greenstones" are similar to the sediments today found in oceanic trenches, above subduction zones. For this reason, greenstones are sometimes seen as evidence for subduction during the Archean. The second type is a complex of felsic magmatic rocks. These rocks are mostly tonalite, trondhjemite or granodiorite, types of rock similar in composition to granite (hence such terranes are called TTG-terranes). TTG-complexes are seen as the relicts of the first continental crust, formed by partial melting in basalt.[60]:Chapter 5

Earth is often described as having had three atmospheres. The first atmosphere, captured from the solar nebula, was composed of light (atmophile) elements from the solar nebula, mostly hydrogen and helium. A combination of the solar wind and Earth's heat would have driven off this atmosphere, as a result of which the atmosphere is now depleted of these elements compared to cosmic abundances.[62] After the impact which created the Moon, the molten Earth released volatile gases; and later more gases were released by volcanoes, completing a second atmosphere rich in greenhouse gases but poor in oxygen. [1]:256 Finally, the third atmosphere, rich in oxygen, emerged when bacteria began to produce oxygen about 2.8 Ga.[63]:83–84, 116–117

In early models for the formation of the atmosphere and ocean, the second atmosphere was formed by outgassing of volatiles from the Earth's interior. Now it is considered likely that many of the volatiles were delivered during accretion by a process known as impact degassing in which incoming bodies vaporize on impact. The ocean and atmosphere would, therefore, have started to form even as the Earth formed.[64] The new atmosphere probably contained water vapor, carbon dioxide, nitrogen, and smaller amounts of other gases.[65]

Planetesimals at a distance of 1 astronomical unit (AU), the distance of the Earth from the Sun, probably did not contribute any water to the Earth because the solar nebula was too hot for ice to form and the hydration of rocks by water vapor would have taken too long.[64]<

In early models for the formation of the atmosphere and ocean, the second atmosphere was formed by outgassing of volatiles from the Earth's interior. Now it is considered likely that many of the volatiles were delivered during accretion by a process known as impact degassing in which incoming bodies vaporize on impact. The ocean and atmosphere would, therefore, have started to form even as the Earth formed.[64] The new atmosphere probably contained water vapor, carbon dioxide, nitrogen, and smaller amounts of other gases.[65]

Planetesimals at a distance of 1 astronomical unit (AU), the distance of the Earth from the Sun, probably did not contribute any water to the Earth because the solar nebula was too hot for ice to form and the hydration of rocks by water vapor would have taken too long.[64][66] The water must have been supplied by meteorites from the outer asteroid belt and some large planetary embryos from beyond 2.5 AU.[64][67] Comets may also have contributed. Though most comets are today in orbits farther away from the Sun than Neptune, computer simulations show that they were originally far more common in the inner parts of the Solar System.[59]:130–132

As the Earth cooled, clouds formed. Rain created the oceans. Recent evidence suggests the oceans may have begun forming as early as 4.4 Ga.[34] By the start of the Archean eon, they already covered much of the Earth. This early formation has been difficult to explain because of a problem known as the faint young Sun paradox. Stars are known to get brighter as they age, and at the time of its formation the Sun would have been emitting only 70% of its current power. Thus, the Sun has become 30% brighter in the last 4.5 billion years.[68] Many models indicate that the Earth would have been covered in ice.[69][64] A likely solution is that there was enough carbon dioxide and methane to produce a greenhouse effect. The carbon dioxide would have been produced by volcanoes and the methane by early microbes. Another greenhouse gas, ammonia, would have been ejected by volcanos but quickly destroyed by ultraviolet radiation.[63]:83

One of the reasons for interest in the early atmosphere and ocean is that they form the conditions under which life first arose. There are many models, but little consensus, on how life emerged from non-living chemicals; chemical systems created in the laboratory fall well short of the minimum complexity for a living organism.[70][71]

The first step in the emergence of life may have been chemical reactions that produced many of the simpler organic compounds, including nucleobases and amino acids, that are the building blocks of life. An experiment in 1953 by Stanley Miller and Harold Urey showed that such molecules could form in an atmosphere of water, methane, ammonia and hydrogen with the aid of sparks to mimic the effect of lightning.[72] Although atmospheric composition was probably different from that used by Miller and Urey, later experiments with more realistic compositions also managed to synthesize organic molecules.[73] Computer simulations show that extraterrestrial organic molecules could have formed in the protoplanetary disk before the formation of the Earth.[74]

Additional complexity could have been reached from at least three possible starting points: self-replication, an organism's ability to produce offspring that are similar to itself; metabolism, its ability to feed and repair itself; and external cell membranes, which allow food to enter and waste products to leave, but exclude unwanted substances.[75]

Replication first: RNA world

Even the simplest members of the three modern domains of life use DNA to record their "recipes" and a complex array of RNA and protein molecules to "read" these instructions and use them for growth, maintenance, and self-replication.

The discovery that a kind of RNA molecule called a ribozyme can catalyze both its own replication and the construction of proteins led to the hypothesis that earlier life-forms were based entirely on RNA.[76] They could have formed an RNA world in which there were individuals but no species, as mutations and horizontal gene transfers would have meant that the offspring in each generation were quite likely to have different genomes from those that their parents started with.[77] RNA would later have been replaced by DNA, which is more stable and therefore can build longer genomes, expanding the range of capabilities a single organism can have.[78] Ribozymes remain as the main components of ribosomes, the "protein factories" of modern cells.[79]

Although short, self-replicating RNA molecules have been artificially produced in laboratories,[80] doubts have been raised about whether natural non-biological synthesis of RNA is possible.[81][82][83] The earliest ribozymes may have been formed of simpler nucleic acids such as PNA, TNA or GNA, which would have been replaced later by RNA.[84][85] Other pre-RNA replicators have been posited, including crystals[86]:150 and even quantum systems.[87]

In 2003 it was proposed that porous metal sulfide precipitates would assist RNA synthesis at about 100 °C (212 °F) and at ocean-bottom pressures near hydrothermal vents. In this hypothesis, the proto-cells would be confined in the pores of the metal substrate until the later development of lipid membranes.[88]

Metabolism first: iron–sulfur world

The replicator in virtually all known life is deoxyribonucleic acid. DNA is far more complex than the original replicator and its replication systems are highly elaborate.

Another long-standing hypothesis is that the first life was composed of protein molecules. Amino acids, the building blocks of proteins, are easily synthesized in plausible prebiotic conditions, as are small peptides (polymers of amino acids) that make good catalysts.[89]:295–297 A series of experiments starting in 1997 showed that amino acids and peptides could form in the presence of carbon monoxide and hydrogen sulfide with iron sulfide and nickel sulfide as catalysts. Most of the steps in their assembly required temperatures of about 100 °C (212 °F) and moderate pressures, although one stage required 250 °C (482 °F) and a pressure equivalent to that found under 7 kilometers (4.3 mi) of rock. Hence, self-sustaining synthesis of proteins could have occurred near hydrothermal vents.[90]

A difficulty with the metabolism-first scenario is finding a way for organisms to evolve. Without the ability to replicate as individuals, aggregates of molecules would have "compositional genomes" (counts of molecular species in the aggregate) as the target of natural selection. However, a recent model shows that such a system is unable to evolve in response to natural selection.[91]

Membranes first: Lipid world

It has been suggested that double-walled "bubbles" of lipids like those that form the external membranes of cells may have been an essential first step.[92] Experiments that simulated the conditions of the early Earth have reported the formation of lipids, and these can spontaneously form liposomes, double-walled "bubbles", and then reproduce themselves. Although they are not intrinsically information-carriers as nucleic acids are, they would be subject to natural selection for longevity and reproduction. Nucleic acids such as RNA might then have formed more easily within the liposomes than they would have outside.[93]

The clay theory

Some clays, notably montmorillonite, have properties that make them plausible accelerators for the emergence of an RNA world: they grow by self-replication of their crystalline pattern, are subject to an analog of natural selection (as the clay "species" that grows fastest in a particular environment rapidly becomes dominant), and can catalyze the formation of RNA molecules.[94] Although this idea has not become the scientific consensus, it still has active supporters.[95]:150–158[86]

Cross-section through a liposome

Research in 2003 reported that montmorillonite could also accelerate the conversion of fatty acids into "bubbles", and that the bubbles could encapsulate RNA attached to the clay. Bubbles can then grow by absorbing additional lipids and dividing. The formation of the earliest cells may have been aided by similar processes.[96]

A similar hypothesis presents self-replicating iron-rich clays as the progenitors of nucleotides, lipids and amino acids.[97]

Last universal ancestor

It is believed that of this multiplicity of protocells, only one line survived. Current phylogenetic evidence suggests that the last universal ancestor (LUA) lived during the early Archean eon, perhaps 3.5 Ga or earlier.[98][99] This LUA cell is the ancestor of all life on Earth today. It was probably a prokaryote, possessing a cell membrane and probably ribosomes, but lacking a nucleus or membrane-bound organelles such as mitochondria or chloroplasts. Like modern cells, it used DNA as its genetic code, RNA for information transfer and protein synthesis, and enzymes to catalyze reactions. Some scientists believe that instead of a single organism being the last universal common ancestor, there were populations of organisms exchanging genes by lateral gene transfer.[98]

Proterozoic Eon

The Proterozoic eon lasted from 2.5 Ga to 542 Ma (million years) ago.[2]:130 In this time span, cratons grew into continents with modern sizes. The change to an oxygen-rich atmosphere was a crucial development. Life developed from prokaryotes into eukaryotes and multicellular forms. The Proterozoic saw a couple of severe ice ages called snowball Earths. After the last Snowball Earth about 600 Ma, the evolution of life on Earth accelerated. About 580 Ma, the Ediacaran biota formed the prelude for the Cambrian Explosion.[citation needed]

Oxygen revolution

Lithified stromatolites on the shores of Lake Thetis, Western Australia. Archean stromatolites are the first direct fossil traces of life on Earth.
A banded iron formation from the 3.15 Ga Moories Group, Barberton Greenstone Belt, South Africa. Red layers represent the times when oxygen was available; gray layers were formed in anoxic circumstances.

The earliest cells absorbed energy and food from the surrounding environment. They used fermentation, the breakdown of more complex compounds into less complex compounds with less energy, and used the energy so liberated to grow and reproduce. Fermentation can only occur in an anaerobic (oxygen-free) environment. The evolution of photosynthesis made it possible for cells to derive energy from the Sun.[100]:377

Most of the life that covers the surface of the Earth depends directly or indirectly on photosynthesis. The most common form, oxygenic photosynthesis, turns carbon dioxide, water, and sunlight into food. It captures the energy of sunlight in energy-rich molecules such as ATP, which then provide the energy to make sugars. To supply the electrons in the circuit, hydrogen is stripped from water, leaving oxygen as a waste product.[101] Some organisms, including purple bacteria and green sulfur bacteria, use an anoxygenic form of photosynthesis that uses alternatives to hydrogen stripped from water as electron donors; examples are hydrogen sulfide, sulfur and iron. Such extremophile organisms are restricted to otherwise inhospitable environments such as hot springs and hydrothermal vents.[100]:379–382[102]

The simpler anoxygenic form arose about 3.8 Ga, not long after the appearance of life. The timing of oxygenic photosynthesis is more controversial; it had certainly appeared by about 2.4 Ga, but some researchers put it back as far as 3.2 Ga.[101] The latter "probably increased global productivity by at least two or three orders of magnitude".[103][104] Among the oldest remnants of oxygen-producing lifeforms are fossil stromatolites.[103][104][61]

At first, the released oxygen was bound up with limestone, iron, and other minerals. The oxidized iron appears as red layers in geological strata called banded iron formations that formed in abundance during the Siderian period (between 2500 Ma and 2300 Ma).[2]:133 When most of the exposed readily reacting minerals were oxidized, oxygen finally began to accumulate in the atmosphere. Though each cell only produced a minute amount of oxygen, the combined metabolism of many cells over a vast time transformed Earth's atmosphere to its current state. This was Earth's third atmosphere.[105]:50–51[63]:83–84, 116–117

Some oxygen was stimulated by solar ultraviolet radiation to form ozone, which collected in a layer near the upper part of the atmosphere. The ozone layer absorbed, and still absorbs, a significant amount of the ultraviolet radiation that once had passed through the atmosphere. It allowed cells to colonize the surface of the ocean and eventually the land: without the ozone layer, ultraviolet radiation bombarding land and sea would have caused unsustainable levels of mutation in exposed cells.[106][59]:219–220

Photosynthesis had another major impact. Oxygen was toxic; much life on Earth probably died out as its levels rose in what is known as the oxygen catastrophe. Resistant forms survived and thrived, and some developed the ability to use oxygen to increase their metabolism and obtain more energy from the same food.[106]

Snowball Earth

The natural evolution of the Sun made it progressively more luminous during the Archean and Proterozoic eons; the Sun's luminosity increases 6% every billion years.[59]:165 As a result, the Earth began to receive more heat from the Sun in the Proterozoic eon. However, the Earth did not get warmer. Instead, the geological record suggests it cooled dramatically during the early Proterozoic. Glacial deposits found in South Africa date back to 2.2 Ga, at which time, based on paleomagnetic evidence, they must have been located near the equator. Thus, this glaciation, known as the Huronian glaciation, may have been global. Some scientists suggest this was so severe that the Earth was frozen over from the poles to the equator, a hypothesis called Snowball Earth.[107]

The Huronian ice age might have been caused by the increased oxygen concentration in the atmosphere, which caused the decrease of methane (CH4) in the atmosphere. Methane is a strong greenhouse gas, but with oxygen it reacts to form CO2, a less effective greenhouse gas.[59]:172 When free oxygen became available in the atmosphere, the concentration of methane could have decreased dramatically, enough to counter the effect of the increasing heat flow from the Sun.[108]

However, the term Snowball Earth is more commonly used to describe later extreme ice ages during the Cryogenian period. There were four periods, each lasting about 10 million years, between 750 and 580 million years ago, when the earth is thought to have been covered with ice apart from the highest mountains, and average temperatures were about −50 °C (−58 °F).[109] The snowball may have been partly due to the location of the supercontinent Rodinia straddling the Equator. Carbon dioxide combines with rain to weather rocks to form carbonic acid, which is then washed out to sea, thus extracting the greenhouse gas from the atmosphere. When the continents are near the poles, the advance of ice covers the rocks, slowing the reduction in carbon dioxide, but in the Cryogenian the weathering of Rodinia was able to continue unchecked until the ice advanced to the tropics. The process may have finally been reversed by the emission of carbon dioxide from volcanoes or the destabilization of methane gas hydrates. According to the alternative Slushball Earth theory, even at the height of the ice ages there was still open water at the Equator.[110][111]

Emergence of eukaryotes

Chloroplasts in the cells of a moss

Modern taxonomy classifies life into three domains. The time of their origin is uncertain. The Bacteria domain probably first split off from the other forms of life (sometimes called Neomura), but this supposition is controversial. Soon after this, by 2 Ga,[112] the Neomura split into the Archaea and the Eukarya. Eukaryotic cells (Eukarya) are larger and more complex than prokaryotic cells (Bacteria and Archaea), and the origin of that complexity is only now becoming known.[citation needed] The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2.4 ago; these multicellular benthic organisms had filamentous structures capable of anastomosis.[113]

Around this time, the first proto-mitochondrion was formed. A bacterial cell related to today's Rickettsia,[114] which had evolved to metabolize oxygen, entered a larger prokaryotic cell, which lacked that capability. Perhaps the large cell attempted to digest the smaller one but failed (possibly due to the evolution of prey defenses). The smaller cell may have tried to parasitize the larger one. In any case, the smaller cell survived inside the larger cell. Using oxygen, it metabolized the larger cell's waste products and derived more energy. Part of this excess energy was returned to the host. The smaller cell replicated inside the larger one. Soon, a stable symbiosis developed between the large cell and the smaller cells inside it. Over time, the host cell acquired some genes from the smaller cells, and the two kinds became dependent on each other: the larger cell could not survive without the energy produced by the smaller ones, and these, in turn, could not survive without the raw materials provided by the larger cell. The whole cell is now considered a single organism, and the smaller cells are classified as organelles called mitochondria.[115]

A similar event occurred with photosynthetic cyanobacteria[116] entering large heterotrophic cells and becoming chloroplasts.[105]:60–61[117]:536–539 Probably as a result of these changes, a line of cells capable of photosynthesis split off from the other eukaryotes more than 1 billion years ago. There were probably several such inclusion events. Besides the well-established endosymbiotic theory of the cellular origin of mitochondria and chloroplasts, there are theories that cells led to peroxisomes, spirochetes led to cilia and flagella, and that perhaps a DNA virus led to the cell nucleus,[118][119] though none of them are widely accepted.[120]

Archaeans, bacteria, and eukaryotes continued to diversify and to become more complex and better adapted to their environments. Each domain repeatedly split into multiple lineages, although little is known about the history of the archaea and bacteria. Around 1.1 Ga, the supercontinent Rodinia was assembling.[121][122] The plant, animal, and fungi lines had split, though they still existed as solitary cells. Some of these lived in colonies, and gradually a division of labor began to take place; for instance, cells on the periphery might have started to assume different roles from those in the interior. Although the division between a colony with specialized cells and a multicellular organism is not always clear, around 1 billion years ago[123], the first multicellular plants emerged, probably green algae.[124] Possibly by around 900 Ma[117]:488 true multicellularity had also evolved in animals.[citation needed]

At first, it probably resembled today's sponges, which have totipotent cells that allow a disrupted organism to reassemble itself.[117]:483–487 As the division of labor was completed in all lines of multicellular organisms, cells became more specialized and more dependent on each other; isolated cells would die.[citation needed]

Supercontinents in the Proterozoic

organic compounds, including nucleobases and amino acids, that are the building blocks of life. An experiment in 1953 by Stanley Miller and Harold Urey showed that such molecules could form in an atmosphere of water, methane, ammonia and hydrogen with the aid of sparks to mimic the effect of lightning.[72] Although atmospheric composition was probably different from that used by Miller and Urey, later experiments with more realistic compositions also managed to synthesize organic molecules.[73] Computer simulations show that extraterrestrial organic molecules could have formed in the protoplanetary disk before the formation of the Earth.[74]

Additional complexity could have been reached from at least three possible starting points: self-replication, an organism's ability to produce offspring that are similar to itself; metabolism, its ability to feed and repair itself; and external cell membranes, which allow food to enter and waste products to leave, but exclude unwanted substances.[75]

Even the simplest members of the three modern domains of life use DNA to record their "recipes" and a complex array of RNA and protein molecules to "read" these instructions and use them for growth, maintenance, and self-replication.

The discovery that a kind of RNA molecule called a ribozyme can catalyze both its own replication and the construction of protei

The discovery that a kind of RNA molecule called a ribozyme can catalyze both its own replication and the construction of proteins led to the hypothesis that earlier life-forms were based entirely on RNA.[76] They could have formed an RNA world in which there were individuals but no species, as mutations and horizontal gene transfers would have meant that the offspring in each generation were quite likely to have different genomes from those that their parents started with.[77] RNA would later have been replaced by DNA, which is more stable and therefore can build longer genomes, expanding the range of capabilities a single organism can have.[78] Ribozymes remain as the main components of ribosomes, the "protein factories" of modern cells.[79]

Although short, self-replicating RNA molecules have been artificially produced in laboratories,[80] doubts have been raised about whether natural non-biological synthesis of RNA is possible.[81][82][83] The earliest ribozymes may have been formed of simpler nucleic acids such as PNA, TNA or GNA, which would have been replaced later by RNA.[84][85] Other pre-RNA replicators have been posited, including crystals[86]:150 and even quantum systems.[87]

In 2003 it was proposed that porous metal sulfide precipitates would assist RNA synthesis at about 100 °C (212 °F) and at ocean-bottom pressures near hydrothermal vents. In this hypothesis, the proto-cells would be confined in the pores of the metal substrate until the later development of lipid membranes.[88]

Another long-standing hypothesis is that the first life was composed of protein molecules. Amino acids, the building blocks of proteins, are easily synthesized in plausible prebiotic conditions, as are small peptides (polymers of amino acids) that make good catalysts.[89]:295–297 A series of experiments starting in 1997 showed that amino acids and peptides could form in the presence of carbon monoxide and hydrogen sulfide with iron sulfide and nickel sulfide as catalysts. Most of the steps in their assembly required temperatures of about 100 °C (212 °F) and moderate pressures, although one stage required 250 °C (482 °F) and a pressure equivalent to that found under 7 kilometers (4.3 mi) of rock. Hence, self-sustaining synthesis of proteins could have occurred near hydrothermal vents.[90]

A difficulty with the metabolism-first scenario is finding a way for organisms to evolve. Without the ability to replicate as individuals, aggregates of molecules would have "compositional genomes" (counts of molecular species in the aggregate) as the target of natural selection. However, a recent model shows that such a system is unable to evolve in response to natural selection.[91]

Membranes first: Lipid world

It has been suggested that double-walled "bubbles" of lipids like those that form the external membranes of cells may have been an essential first step.[92] Experiments that simulated the conditions of the early Earth have reported the formation of lipids, and these can spontaneously form liposomes, double-walled "bubbles", and then reproduce themselves. Although they are not intrinsically information-carriers as nucleic acids are, they would be subject to natural selection for longevity and reproduction. Nucleic acids such as RNA might then have formed more easily within the liposo

A difficulty with the metabolism-first scenario is finding a way for organisms to evolve. Without the ability to replicate as individuals, aggregates of molecules would have "compositional genomes" (counts of molecular species in the aggregate) as the target of natural selection. However, a recent model shows that such a system is unable to evolve in response to natural selection.[91]

It has been suggested that double-walled "bubbles" of lipids like those that form the external membranes of cells may have been an essential first step.[92] Experiments that simulated the conditions of the early Earth have reported the formation of lipids, and these can spontaneously form liposomes, double-walled "bubbles", and then reproduce themselves. Although they are not intrinsically information-carriers as nucleic acids are, they would be subject to natural selection for longevity and reproduction. Nucleic acids such as RNA might then have formed more easily within the liposomes than they would have outside.[93]

The clay theory

clays, notably montmorillonite, have properties that make them plausible accelerators for the emergence of an RNA world: they grow by self-replication of their crystalline pattern, are subject to an analog of natural selection (as the clay "species" that grows fastest in a particular environment rapidly becomes dominant), and can catalyze the formation of RNA molecules.[94] Although this idea has not become the scientific consensus, it still has active supporters.[95]:150–158[86]

Research in 2003 reported that montmorillonite could also accelerate the conversion of fatty acids into "bubbles", and that the bubbles could encapsulate RNA attached to the clay. Bubbles can then grow by absorbing additional lipids and dividing. The formation of the earliest cells may have been aided by similar processes.[96]

A similar hypothesis presents self-replicating iron-rich clays as the progenitors of nucleotides, lipids and amino acids.[97]

Last universal ancestor

It is believed that of this multiplicity of protocells, only one nucleotides, lipids and amino acids.[97]

It is believed that of this multiplicity of protocells, only one line survived. Current phylogenetic evidence suggests that the last universal ancestor (LUA) lived during the early Archean eon, perhaps 3.5 Ga or earlier.[98][99] This LUA cell is the ancestor of all life on Earth today. It was probably a prokaryote, possessing a cell membrane and probably ribosomes, but lacking a nucleus or membrane-bound organelles such as mitochondria or chloroplasts. Like modern cells, it used DNA as its genetic code, RNA for information transfer and protein synthesis, and enzymes to catalyze reactions. Some scientists believe that instead of a single organism being the last universal common ancestor, there were populations of organisms exchanging genes by lateral gene transfer.[98]

Proterozoic Eon

The Proterozoic eon lasted from 2.5 Ga to 542 Ma (million y

The Proterozoic eon lasted from 2.5 Ga to 542 Ma (million years) ago.[2]:130 In this time span, cratons grew into continents with modern sizes. The change to an oxygen-rich atmosphere was a crucial development. Life developed from prokaryotes into eukaryotes and multicellular forms. The Proterozoic saw a couple of severe ice ages called snowball Earths. After the last Snowball Earth about 600 Ma, the evolution of life on Earth accelerated. About 580 Ma, the Ediacaran biota formed the prelude for the Cambrian Explosion.[citation needed]

Oxygen revolution

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