Skip to content
— CH. 1 · THE FIRST LIVING THING —

Abiogenesis

~11 min read · Ch. 1 of 8
8 sections
  • NASA defines life with a single tight phrase: "a self-sustaining chemical system capable of Darwinian evolution." That definition sounds simple until you try to build one from rock and water. Abiogenesis, sometimes called biopoiesis, is the natural process by which life arises from non-living matter, such as simple organic compounds. The transition from non-life to life has never been observed in any experiment. Researchers can only propose stages for how it happened.

    Consider a problem that sits at the heart of the whole inquiry. A cell copies its DNA using the DNA polymerase enzyme. That enzyme is itself produced by translating the DNA polymerase gene. Neither the enzyme nor the DNA can be made without the other. So how does a system this tightly interlocked assemble itself one step at a time, when every part seems to need every other part to work at all?

    The last universal common ancestor of all modern life, known as LUCA, lived some 4 billion years ago. It was already a complex single-celled organism with hundreds of genes. It was not the first living thing. Between the true origin of life and LUCA lies an unknown span of gene transfers, extinctions, and adaptation. The chapters ahead trace how chemistry on a young planet might have crossed that gap, from the first stars that forged carbon to the warm pools and deep vents where life may have begun.

  • On the 1st of February 1871, Charles Darwin wrote to Joseph Hooker and let himself speculate. The original spark of life, he mused, may have formed in a "warm little pond, with all sorts of ammonia and phosphoric salts,-light, heat, electricity present, that a protein compound was chemically formed." Darwin added a sharp observation: such matter today would be "instantly devoured or absorbed," which would not have been the case before living creatures existed.

    Long before Darwin, the dominant view was spontaneous generation. From Aristotle until the 19th century, people believed lower animals like insects sprang from decaying organic matter. In 1665, Robert Hooke published the first drawings of a microorganism. In 1676, Antonie van Leeuwenhoek drew and described microorganisms, probably protozoa and bacteria, and by the 1680s convinced himself the old theory was wrong. In 1668, Francesco Redi showed that no maggots appeared in meat when flies were kept from laying eggs.

    Alexander Oparin in 1924 and J. B. S. Haldane in 1929 turned the pond into a theory. The earliest cells, they proposed, slowly self-organized from a primordial soup. Haldane pictured prebiotic oceans as a "hot dilute soup" in which organic compounds could form. In 1967, J. D. Bernal laid out three stages: the origin of biological monomers, the origin of biological polymers, and the evolution from molecules to cells.

    A rival idea, panspermia, dates back to Anaxagoras in the 5th century BC. It holds that life originated elsewhere in the universe and arrived here, perhaps carried by meteoroids, asteroids, or comets. There remains some interest in the possibility that life began on Mars and later transferred to Earth.

  • In 1952, Stanley Miller and Harold Urey sealed a highly reducing mixture of gases, methane, ammonia, and hydrogen, together with water vapor, and ran it through a chemical experiment. Out came organic monomers, including amino acids, formed from inorganic precursors alone. It was a demonstration that the building blocks of life could appear spontaneously under conditions like those of early Earth.

    Bernal was not fully satisfied. He said it was not enough to explain how such molecules form. What was needed, he argued, was a physical-chemical account of "suitable sources and sinks for free energy." Later science complicated the picture further. The primitive atmosphere is now described as weakly reducing or neutral, which shrinks the variety of amino acids the original setup could yield. Adding iron and carbonate minerals, present in early oceans, restores a diverse array of amino acids.

    Alexander Butlerov showed in 1861 that the formose reaction creates sugars, including tetroses, pentoses, and hexoses, when formaldehyde is heated under basic conditions with divalent metal ions like calcium. R. Breslow proposed in 1959 that the reaction was autocatalytic. Nucleobases such as guanine and adenine can be built from simple sources like hydrogen cyanide and ammonia. Formamide, made from water and HCN, produces all four ribonucleotides when warmed with terrestrial minerals.

    Cold proves surprisingly productive. Freezing temperatures assist the synthesis of purines by concentrating precursors like HCN. In one striking result, seven amino acids and eleven types of nucleobases formed in ice when ammonia and cyanide were left in a freezer for 25 years. These low-temperature reactions run fast because eutectic freezing crowds impurities into microscopic pockets of liquid within the ice.

  • During the Late Heavy Bombardment, meteorites may have delivered up to five million tons of organic prebiotic material to Earth each year. Even now, 40,000 tons of cosmic dust fall to Earth annually. Space, it turns out, is a chemistry set of its own. Organic compounds are relatively common there, built in "factories of complex molecular synthesis" within molecular clouds and circumstellar envelopes.

    The amino acid glycine was found in material ejected from comet Wild 2, having earlier been detected in meteorites. Purine and pyrimidine nucleobases, including guanine, adenine, cytosine, uracil, and thymine, along with sugars, have turned up in meteorites. NASA studies suggest all four DNA nucleobases formed in outer space. Glycolaldehyde, a sugar molecule and RNA precursor, has been detected around protostars and on meteorites.

    Polycyclic aromatic hydrocarbons, abbreviated PAHs, are the most common and abundant polyatomic molecules in the observable universe and a major store of carbon. They seem to have formed shortly after the Big Bang and are associated with new stars and exoplanets. A star called HH 46-IR, resembling the early Sun, is wrapped in a disk containing cyanide compounds, hydrocarbons, and carbon monoxide.

    Carbon itself has a stellar pedigree. It formed mainly in white dwarf stars, which ejected carbon and oxygen across the universe as they died, allowing rocky planets to form. Soon after the Big Bang, roughly 14 Gya, only hydrogen, helium, and lithium existed. The first stars fused hydrogen and forged heavier elements up to Iron-56, with the heaviest born in supernovas. The cosmic dust permeating the universe still carries complex organics, amorphous solids with a mixed aromatic-aliphatic structure, that stars can create rapidly.

  • The age of the Earth is 4.54 Gya, found by radiometric dating of calcium-aluminium-rich inclusions in carbonaceous chondrite meteorites, the oldest material in the Solar System. Soon after initial accretion at 4.48 Gya, a collision with a hypothesised impactor named Theia is thought to have thrown off the debris that became the Moon. That impact stripped away Earth's primary atmosphere.

    What remained was an atmosphere largely of water vapor, nitrogen, and carbon dioxide, with smaller amounts of carbon monoxide, hydrogen, and sulfur compounds. Carbon dioxide dissolving in water made the early seas slightly acidic, around a pH of 5.5. Liquid oceans may have condensed as early as the Moon-forming impact. That scenario draws support from 4.404 Gya zircon crystals with high oxygen-18 values, recovered from metamorphosed quartzite at Mount Narryer in Western Australia.

    The Hadean atmosphere has been called a "gigantic, productive outdoor chemical laboratory." Earth may have been a predominantly water world between 4.4 and 4.3 Gya, swept by intense ultraviolet light from a Sun in its T Tauri stage and battered by asteroid and comet impacts. Whether any crust rose above that ocean is still debated.

    The Late Heavy Bombardment hypothesis once imagined a single cataclysmic strike at 3.9 Gya that could have sterilized the planet by boiling its oceans. That 3.9 Gya date came from Apollo samples gathered mostly near the Imbrium Basin, which biased the record. Impact modelling now suggests many small, short-lived bombardment periods instead, with terrestrial ejecta layers appearing both before and after the 3.9 Gya marker. A young Earth under continuous lighter impacts had far less power to wipe the slate clean.

  • In 2017, the earliest physical evidence of life was reported as microbialites in the Nuvvuagittuq Greenstone Belt of Northern Quebec, set in banded iron formation rocks at least 3.77 and possibly as old as 4.32 Gya. The micro-organisms could have lived within hydrothermal vent precipitates, soon after oceans formed around 4.4 Gya. They resemble modern hydrothermal vent bacteria. Later research disputed the reading, arguing abiotic processes in silica-rich waters or volcanic ejecta might explain the same observations.

    Biogenic graphite appears in 3.7 Gya metasedimentary rocks from southwestern Greenland. Near the Isua supracrustal belt, rocks from Akilia Island dating to 3.7 Gya show biogenic carbon isotopes. Within the Isua belt, graphite inclusions trapped in garnet crystals connect to oxygen, nitrogen, and possibly phosphorus. Carbon isotope ratios on graphite inclusions from the Jack Hills zircons suggest life could have existed from 4.1 Gya.

    The Pilbara region of Western Australia holds the Dresser Formation, with rocks 3.48 Gya containing layered structures called stromatolites. Their modern counterparts are made by photosynthetic micro-organisms, including cyanobacteria. Parts of the formation preserve hot springs on land, while other regions appear to have been shallow seas. In the same region, pyrite-bearing sandstone from a fossilized beach held rounded tubular cells that oxidized sulfur by photosynthesis without oxygen.

    A 2024 study inferred LUCA's age as around 4.2 Gya, within a range of 4.09 to 4.33 Gya, by analysing pre-LUCA gene duplicates. That places LUCA much sooner after the origin of life than previously thought. A molecular clock analysis suggests LUCA emerged prior to 3.9 Gya, hinting that life took hold quickly in geological terms.

  • The largest unanswered question in evolution is how simple protocells first arose and then differed in their contribution to the next generation, setting evolution in motion. The lipid world theory proposes that the first self-replicating object was lipid-like. Phospholipids form lipid bilayers in water under agitation, but those molecules were not present on early Earth. Other membrane-forming amphiphilic long-chain molecules were, and such bodies can expand by inserting lipids and split into two offspring of similar size.

    Irene Chen and Jack W. Szostak suggest that elementary protocells can give rise to primitive differential reproduction, competition, and energy storage. Competition for membrane molecules would favor stabilized membranes, hinting at a selective advantage for cross-linked fatty acids and even modern phospholipids. A membrane is exactly what a cell needs to create its own electrochemical gradient. Fatty acid vesicles in alkaline vent conditions can be stabilized by isoprenoids made through the formose reaction.

    In 1961, Peter Mitchell proposed chemiosmosis as a cell's primary system of energy conversion. The mechanism now powers energy conversion in micro-organisms and in the mitochondria of eukaryotes, which produce adenosine triphosphate, or ATP, the energy currency of the cell. ATP synthesis runs on protons moving across a closed membrane where the ATP synthase enzyme sits. In the first organisms, that proton gradient could have come from the difference between flow out of a hydrothermal vent and the surrounding seawater.

    Life requires a loss of entropy as molecules organize into living matter. That does not break the second law of thermodynamics. A living organism creates order in its body at the expense of more entropy elsewhere, in heat and waste. A boundary is therefore needed to separate ordered life from chaotic non-living matter, and a functional protocell capable of Darwinian evolution has not yet been built in any laboratory.

  • William Martin and Michael Russell proposed that life began in metal-sulphide-walled compartments at submarine hydrothermal vents, structures that could act as precursors to cell walls. These vents form where hydrogen-rich fluids rise from below the sea floor through serpentinization of olivine, meeting carbon-dioxide-rich ocean water at a pH interface. The redox reactions there, with molecular hydrogen meeting carbon dioxide, supply a sustained chemical energy source. Russell showed that alkaline vents create an abiogenic proton motive force, a chemiosmotic gradient he called ideal for abiogenesis.

    Not everyone is convinced the vents are the cradle. Vents do not concentrate prebiotic materials, since strong dilution by seawater leaves little residence time to accumulate compounds. All modern cells rely on phosphates for the nucleotide backbone and potassium for protein formation, yet both were scarce in Archaean oceans. No studies have experimentally demonstrated the synthesis of sugars, amino acids, nucleotides, or membrane-forming fatty acids under plausible vent conditions.

    Surface bodies of water offer something the open ocean cannot: wet-dry cycles that concentrate prebiotic compounds and enable condensation reactions to build macromolecules. Lakes and ponds also receive phosphate from weathering of continental apatite-containing rocks. Mulkidjanian and co-authors argue that only hot springs, like those at Kamchatka, provide the ionic balance cells universally need, especially a high potassium to sodium ratio. A phylogenomic analysis found that LUCA's intracellular fluid matches the ionic composition of hot springs.

    Cold-start theories take the opposite tack. Stellar models predict the early Sun was about 25% weaker than today, which would favor an icy planet, though strong evidence points to liquid water alongside icy poles. Water under ice would be shielded from ultraviolet light while staying connected through cracks. Experiments simulating Europa-like conditions near 20 degrees Celsius have synthesised amino acids and adenine.

    There is also a case for going underground. The continental crust, threaded with tectonic fault zones, could offer a stable, well-protected setting for long-term prebiotic evolution. At depths around 1000 meters, carbon dioxide sits near the phase transition between supercritical and gaseous states, letting lipophilic molecules accumulate and precipitate. One quirk of life narrows the search further. Living organisms are homochiral, with amino acids almost always left-handed and sugars right-handed, and some amino acids delivered by meteorites already show a chiral asymmetry of the same sign as life on Earth.

Common questions

What is abiogenesis and how does it explain the origin of life?

Abiogenesis, sometimes called biopoiesis, is the natural process by which life arises from non-living matter such as simple organic compounds. The prevailing scientific view is that it was not a single event but a process of increasing complexity, including the prebiotic synthesis of organic molecules, molecular self-replication, self-assembly, autocatalysis, and the emergence of cell membranes.

When did life first appear on Earth according to abiogenesis research?

Based on the geologic record, life most likely emerged on Earth between 4.32 and 3.48 Gya. The earliest physical evidence reported in 2017 consists of microbialites in the Nuvvuagittuq Greenstone Belt of Northern Quebec, in rocks at least 3.77 and possibly as old as 4.32 Gya. The Earth itself formed at 4.54 Gya.

What did the Miller-Urey experiment prove about the origin of life?

In 1952, Stanley Miller and Harold Urey showed that organic monomers such as amino acids could form spontaneously from inorganic precursors under prebiotic conditions. They used a highly reducing mixture of methane, ammonia, and hydrogen with water vapor. Later work found that current consensus describes the primitive atmosphere as weakly reducing or neutral, which reduces the variety of amino acids produced.

What is the RNA world hypothesis in abiogenesis?

The RNA world hypothesis describes an early Earth with self-replicating and catalytic RNA but no DNA or proteins. It was proposed in 1962 by Alexander Rich, and the term was coined by Walter Gilbert in 1986. Many researchers agree an RNA world must have preceded modern DNA-based life, though it may not have been the first system to exist.

What was LUCA in the study of the origin of life?

LUCA is the last universal common ancestor of all modern life-forms, a complex single-celled organism that lived over 4 Gya with hundreds of genes. A 2016 study identified 355 genes likely present in LUCA, suggesting it was anaerobic with a Wood-Ljungdahl pathway, nitrogen and carbon fixing, and thermophilic. A 2024 study inferred LUCA's age as around 4.2 Gya.

Where did life on Earth begin according to abiogenesis theories?

Several geological settings have been proposed, often in competition. The deep sea hydrothermal vent theory, advanced by William Martin and Michael Russell, places life at alkaline submarine vents, while other hypotheses favor surface bodies of water, hot springs similar to those at Kamchatka, icy environments, or inside the continental crust along tectonic fault zones.

Did the building blocks of life come from space in abiogenesis?

Many organic building blocks have been found beyond Earth. The amino acid glycine was found in material ejected from comet Wild 2, and nucleobases including guanine, adenine, cytosine, uracil, and thymine have been found in meteorites. During the Late Heavy Bombardment, meteorites may have delivered up to five million tons of organic prebiotic material to Earth per year.

All sources

279 references cited across the entry

  1. 1journalVocabulary of Definitions of Life Suggests a DefinitionEdward N. Trifonov — 17 March 2011
  2. 2webAbout Life DetectionMary A. Voytek — NASA — 6 March 2021
  3. 3bookThe Origin of LifeAleksandr Ivanovich Oparin — Courier — 2003
  4. 4journalA Strategy for Origins of Life ResearchCaleb Scharf — 18 December 2015
  5. 7bookEncyclopedia of AstrobiologyStephane Tirard — 20 April 2015
  6. 8bookThe Emergence of Life: From Chemical Origins to Synthetic BiologyPier Luigi Luisi — Cambridge University Press — 2018
  7. 9journalThe origin of life: what we know, what we can know and what we will never knowAddy Pross et al. — 2013
  8. 10journalRe-conceptualizing the origins of lifeSara I. Walker et al. — 13 November 2017
  9. 11journalExtraterrestrial Life in the UniverseRobert W. Graham — February 1990
  10. 12harvnbAltermann (2009) p. xviiAltermann — 2009
  11. 14harvnbOparin (1953) p. viOparin — 1953
  12. 15journalControversies on the origin of lifeJuli Peretó — 2005
  13. 16journalDid Life Come from Another World?David Warmflash et al. — November 2005
  14. 17harvnbYarus (2010) p. 47Yarus — 2010
  15. 18bookA New History of Life: the radical discoveries about the origins and evolution of life on earthPeter Ward et al. — Bloomsbury Press — 2015
  16. 19harvnbSheldon (2005)Sheldon — 2005
  17. 20harvnbLennox (2001) p. 229–258Lennox — 2001
  18. 21harvnbBernal (1967)Bernal — 1967
  19. 22journalDevelopment of Biology in Aristotle and Theophrastus: Theory of Spontaneous GenerationD. M. Balme — 1962
  20. 23harvnbRoss (1652)Ross — 1652
  21. 24harvnbDobell (1960)Dobell — 1960
  22. 25harvnbBondeson (1999)Bondeson — 1999
  23. 27harvnbOparin (1953) p. 196Oparin — 1953
  24. 28harvnbTyndall (1905) p. IV, XII (1876), XIII (1878)Tyndall — 1905
  25. 29journalSpace MicrobiologyGerda Horneck et al. — March 2010
  26. 30journalBacterial morphologies supporting cometary panspermia: a reappraisalChandra Wickramasinghe — 2011
  27. 31newsVisions of Life on Mars in Earth's DepthsKenneth Chang — 12 September 2016
  28. 34webOrigin and Evolution of Life on a Frozen EarthJohn C. Priscu — National Science Foundation
  29. 35newsCharles Darwin's hunch about early life was probably rightMichael Marshall — 11 November 2020
  30. 37journalPhotochemical Formation of Self-Sustaining CoacervatesKrishna Bahadur — 1975
  31. 38harvnbBryson (2004) p. 300–302Bryson — 2004
  32. 39harvnbBernal (1951)Bernal — 1951
  33. 40journalThe Physical Basis of LifeJohn Desmond Bernal — September 1949
  34. 42journalPrimordial synthesis of amines and amino acids in a 1958 Miller H2S-rich spark discharge experimentEric T. Parker et al. — 5 April 2011
  35. 43harvnbBernal (1967) p. 143Bernal — 1967
  36. 44journalA Reassessment of Prebiotic Organic Synthesis in Neutral Planetary AtmospheresH. James Cleaves et al. — April 2008
  37. 45journalRethinking Earth's Early AtmosphereChristopher F. Chyba — 13 May 2005
  38. 46harvnbBarton, Briggs, Eisen (2007) p. 93–95Barton, Briggs, Eisen — 2007
  39. 47harvnbBada, Lazcano (2009) p. 56–57Bada, Lazcano — 2009
  40. 48journalPrebiotic Soup – Revisiting the Miller ExperimentJeffrey L. Bada et al. — 2 May 2003
  41. 49journalCosmic Star-Formation HistoryPiero Madau et al. — 2014-08-18
  42. 50journalCarbon star formation as seen through the non-monotonic initial–final mass relationPaola Marigo — 6 July 2020
  43. 52webFormation of Solar Systems: Solar Nebular TheoryUniversity of Massachusetts Amherst
  44. 53webAge of the EarthUnited States Geological Survey — 9 July 2007
  45. 54harvnbDalrymple (2001) p. 205–221Dalrymple — 2001
  46. 55harvnbFesenkov (1959) p. 9Fesenkov — 1959
  47. 56journalDating the Moon-forming impact event with asteroidal meteoritesW. F. Bottke et al. — 2015-04-17
  48. 57journalEarth's Early AtmosphereJames F. Kasting — 12 February 1993
  49. 58journalDarwin's warm little pond revisited: from molecules to the origin of lifeHartmut Follmann et al. — November 2009
  50. 59journalHadean Ocean Carbonate GeochemistryJohn Morse — September 1998
  51. 60journalTerrestrial aftermath of the Moon-forming impactNorman H. Sleep et al. — 2014-09-13
  52. 61journalHadean Ocean Carbonate GeochemistryJohn W. Morse et al. — 1998
  53. 62journalDetrital Zircon from the Jack Hills and Mount Narryer, Western Australia: Evidence for Diverse >4.0 Ga Source RocksJames L. Crowley et al. — May 2005
  54. 63journalEvidence from detrital zircons for the existence of continental crust and oceans on the Earth 4.4 Gyr agoSimon A. Wilde et al. — 11 January 2001
  55. 64journalPlate tectonics, flood basalts and the evolution of Earth's oceansJun Korenaga — December 2008
  56. 66journalIsotopic evidence for a terminal lunar cataclysmFouad Tera et al. — April 1974
  57. 67journalCratering History and Lunar ChronologyD. Stoffler — 2006-01-01
  58. 68journalAnnihilation of ecosystems by large asteroid impacts on the early EarthNorman H. Sleep et al. — December 1989
  59. 69journalImpact bombardment of the terrestrial planets and the early history of the Solar SystemCaleb I. Fassett et al. — 2013-06-23
  60. 70journalMicrobial habitability of the Hadean Earth during the late heavy bombardmentOleg Abramov et al. — May 2009
  61. 71journalIllusory Late Heavy BombardmentsPatrick Boehnke et al. — 2016-09-12
  62. 72journalCataclysm No More: New Views on the Timing and Delivery of Lunar ImpactorsNicolle E. B. Zellner — 2017-05-03
  63. 73journalThe terrestrial record of Late Heavy BombardmentDonald R. Lowe et al. — 2018-04-01
  64. 74harvnbBock, Goode (1996)Bock, Goode — 1996
  65. 75journalEvidence for early life in Earth's oldest hydrothermal vent precipitatesMatthew S. Dodd et al. — 1 March 2017
  66. 76journalBiomimetic mineral self-organization from silica-rich spring watersJuan Manuel García-Ruiz et al. — 2017-03-03
  67. 77journalEarth's earliest and deepest purported fossils may be iron-mineralized chemical gardensSean McMahon — 2019-12-04
  68. 78journalOn the biogenicity of Fe-oxyhydroxide filaments in silicified low-temperature hydrothermal deposits: Implications for the identification of Fe-oxidizing bacteria in the rock recordKaren C. Johannessen et al. — January 2020
  69. 79journalRemarkably preserved tephra from the 3430 Ma Strelley Pool Formation, Western Australia: Implications for the interpretation of Precambrian microfossilsDavid Wacey et al. — April 2018
  70. 80journalEvidence for biogenic graphite in early Archaean Isua metasedimentary rocksYoko Ohtomo et al. — January 2014
  71. 81journalMicrobially Induced Sedimentary Structures Recording an Ancient Ecosystem in the ca. 3.48 Gyo Dresser Formation, Pilbara, Western AustraliaNora Noffke et al. — 16 November 2013
  72. 82harvnbDavies (1999)Davies — 1999
  73. 83journalElements of Eoarchean life trapped in mineral inclusionsT. Hassenkam et al. — 2017
  74. 84journalOldest reliable fossils show early life was a beachJames O'Donoghue — 21 August 2011
  75. 85journalMicrofossils of sulphur-metabolizing cells in 3.4-billion-year-old rocks of Western AustraliaDavid Wacey et al. — October 2011
  76. 86journalPotentially biogenic carbon preserved in a 4.1 billion-year-old zirconElizabeth A. Bell et al. — 2015-11-24
  77. 87journalThe nature of the last universal common ancestor and its impact on the early Earth systemEdmund R. R. Moody et al. — 12 July 2024
  78. 89journalEarliest signs of life on land preserved in ca. 3.5 Gao hot spring depositsTara Djokic et al. — 9 May 2017
  79. 90journalIntegrated genomic and fossil evidence illuminates life's early evolution and eukaryote originHolly C. Betts et al. — 20 August 2018
  80. 91webBuilding Blocks of Life's Building Blocks Come From StarlightElizabeth Landau — 12 October 2016
  81. 92journalInfrared diffuse interstellar bands in the Galactic Centre regionThomas R. Geballe et al. — 10 November 2011
  82. 93harvnbKlyce (2001)Klyce — 2001
  83. 94bookOrigins of Life: A Cosmic PerspectiveDouglas Whittet — IOP Publishing — 2017
  84. 96journalCosmic carbon chemistry: from the interstellar medium to the early Earth.Pascale Ehrenfreund et al. — December 2010
  85. 97journalWater and carbon dioxide as the main precursors of organic matter on Earth and in spaceVladislav V. Goncharuk et al. — February 2015
  86. 99journalIdentifying the wide diversity of extraterrestrial purine and pyrimidine nucleobases in carbonaceous meteoritesYasuhiro Oba — 26 April 2022
  87. 101journalColoration and darkening of methane clathrate and other ices by charged particle irradiation: Applications to the outer solar systemWilliam Reid Thompson et al. — 30 December 1987
  88. 102journalPrebiotic Chemistry within a Simple Impacting Icy MixtureNir Goldman et al. — 20 June 2013
  89. 105journalVariations in the Peak Position of the 6.2 μm Interstellar Emission Feature: A Tracer of N in the Interstellar Polycyclic Aromatic Hydrocarbon PopulationDouglas M. Hudgins et al. — 10 October 2005
  90. 106webCosmic Distribution of Chemical ComplexityDavid J. Des Marais et al. — NASA — 2009
  91. 107newsLife's Building Blocks 'Abundant in Space'Bjorn Carey — Imaginova — 18 October 2005
  92. 108journalFormation of Fullerenes in H-Containing Planetary NebulaeDomingo. A. García-Hernández et al. — 20 November 2010
  93. 109journalIn-situ Probing of Radiation-induced Processing of Organics in Astrophysical Ice Analogs – Novel Laser Desorption Laser Ionization Time-of-flight Mass Spectroscopic StudiesMurthy S. Gudipati et al. — 1 September 2012
  94. 110journalAstrochemistry and the origin of genetic materialEnzo Gallori — June 2011
  95. 111journalOrganic Chemistry of Carbonaceous MeteoritesZita Martins — February 2011
  96. 112journalExtraterrestrial nucleobases in the Murchison meteoriteZita Martins et al. — 15 June 2008
  97. 113journalCarbonaceous meteorites contain a wide range of extraterrestrial nucleobasesMichael P. Callahan et al. — 23 August 2011
  98. 114webNASA Researchers: DNA Building Blocks Can Be Made in SpaceJohn Steigerwald — NASA — 8 August 2011
  99. 115journalMixed aromatic–aliphatic organic nanoparticles as carriers of unidentified infrared emission featuresSun Kwok et al. — 3 November 2011
  100. 117journalExtraterrestrial ribose and other sugars in primitive meteoritesYoshihiro Furukawa et al. — 2019-11-13
  101. 118journalSynthesis of purines under possible primitive earth conditions: II. Purine intermediates from hydrogen cyanideJoan Oró et al. — February 1962
  102. 119journalThe origin of the biologically coded amino acidsHenderson Cleaves II — 2010
  103. 120journalPrebiotic Synthesis of N-Formylaminonitriles and Derivatives in FormamideNicholas J. Green et al. — 2023
  104. 121journalOn the Mechanism of the Formose ReactionR. Breslow — 1959
  105. 122journalMechanism of Synthesis of Adenine from Hydrogen Cyanide under Possible Primitive Earth ConditionsJoan Oró — 16 September 1961
  106. 123journalOrigin-of-life Molecules in the Atmosphere after Big Impacts on the Early EarthNicholas F. Wogan et al. — 2023-09-01
  107. 124journalFormamide and the origin of life.Raffaele Saladino et al. — March 2012
  108. 125journalFrom the one-carbon amide formamide to RNA all the steps are prebiotically possibleRaffaele Saladino et al. — July 2012
  109. 126journalSynthesis of activated pyrimidine ribonucleotides in prebiotically plausible conditionsMatthew W. Powner et al. — May 2009
  110. 128journalHigh-energy chemistry of formamide: A unified mechanism of nucleobase formationMartin Ferus et al. — 2015
  111. 129journalTowards a prebiotic chemoton – nucleotide precursor synthesis driven by the autocatalytic formose reactionQuoc Phuong Tran et al. — 2023-09-13
  112. 130journalPrebiotic syntheses of purines and pyrimidinesBrenda Basile et al. — 1984
  113. 131journalPrebiotic Adenine Revisited: Eutectics and PhotochemistryLeslie E. Orgel — August 2004
  114. 132journalAn efficient prebiotic synthesis of cytosine and uracilMichael P. Robertson et al. — 29 June 1995
  115. 133journalDid Life Evolve in Ice?Douglas Fox — 1 February 2008
  116. 134journalPrebiotic Synthesis of Adenine and Amino Acids Under Europa-like ConditionsMatthew Levy et al. — June 2000
  117. 135journalSynthesis of Pyrimidines and Triazines in Ice: Implications for the Prebiotic Chemistry of NucleobasesCésar Menor-Salván et al. — 20 April 2009
  118. 136journalChemical evolution: The mechanism of the formation of adenine under prebiotic conditionsDebjani Roy et al. — 30 October 2007
  119. 137journalPrebiotic Peptides: Molecular Hubs in the Origin of LifeMoran Frenkel-Pinter et al. — 2020-06-10
  120. 138journalPrebiotic condensation through wet–dry cycling regulated by deliquescenceThomas D. Campbell et al. — 2019
  121. 139journalCatalytic peptide hydrolysis by mineral surface: Implications for prebiotic chemistryKarina Marshall-Bowman et al. — October 2010
  122. 140journalHeat flows enrich prebiotic building blocks and enhance their reactivityThomas Matreux et al. — April 2024
  123. 141journalThe possible role of clays in prebiotic peptide synthesisMella Paecht-Horowitz — 1974-01-01
  124. 142journalA pathway to peptides in space through the condensation of atomic carbonS. A. Krasnokutski et al. — 2022-02-10
  125. 143journalFormation of extraterrestrial peptides and their derivativesSerge A. Krasnokutski et al. — 2024-04-19
  126. 144webSystems Prebiology-Studies of the origin of LifeDoron Lancet — Department of Molecular Genetics; Weizmann Institute of Science — 30 December 2014
  127. 145journalThe Lipid WorldDaniel Segré et al. — February 2001
  128. 147journalExcess Mutual Catalysis Is Required for Effective EvolvabilityOmer Markovitch et al. — Summer 2012
  129. 148journalOrigin of Evolution versus Origin of Life: A Shift of ParadigmMarc Tessera — 2011
  130. 149journalFrom Self-Assembled Vesicles to ProtocellsIrene A. Chen et al. — July 2010
  131. 150journalReciprocal Relations in Irreversible Processes ILars Onsager — 1931
  132. 151journalReciprocal Relations in Irreversible Processes IILars Onsager — 1931
  133. 152bookAn Introduction to the Thermodynamics of Irreversible ProcessesIlya Prigogine — Wiley — 1967
  134. 153journalA Simpler Origin for LifeRobert Shapiro — June 2007
  135. 154journalProgress in synthesizing protocellsO. Duhan Toparlak et al. — March 2019
  136. 155webProtocellsNational Science Foundation
  137. 156journalThe Emergence of Cells During the Origin of LifeIrene A. Chen — 8 December 2006
  138. 157magazineWhat Came Before DNA?Carl Zimmer — 26 June 2004
  139. 158harvnbChang (2007)Chang — 2007
  140. 159journalIsoprenoids enhance the stability of fatty acid membranes at the emergence of life potentially leading to an early lipid divideSean F. Jordan et al. — 2019-12-06
  141. 160journalMolecular Evolution in a Peptide-Vesicle SystemChristian Mayer et al. — 2018
  142. 161journalPeriodic Vesicle Formation in Tectonic Fault Zones—an Ideal Scenario for Molecular EvolutionChristian Mayer et al. — 2015-06-01
  143. 162journalMembrane Structure Obtained in an Experimental Evolution ProcessMaría J. Dávila et al. — 2022
  144. 163journalSpontaneous Formation of Functional Structures in Messy EnvironmentsChristian Mayer — 2022
  145. 164journalStructural Phenomena in a Vesicle Membrane Obtained through an Evolution Experiment: A Study Based on MD SimulationsMaría J. Dávila et al. — 2023
  146. 165journalLife in The Context of Order and ComplexityChristian Mayer — 2020-01-18
  147. 166bookHabitability of the Universe Before Earth: Life Before EarthAlexei A. Sharov et al. — Academic Press — 2018
  148. 167journalWhat is a Complex System?J. Ladyman et al. — 2013
  149. 168journalEntropy production as correlation between system and reservoirM. Esposito et al. — 2010
  150. 169journalShock Synthesis of Amino Acids in Simulated Primitive EnvironmentsA. Bar-Nun et al. — 24 April 1970
  151. 170journalCavitation during Impact of Liquid Water on Water: Geochemical ImplicationsMichael Anbar — 27 September 1968
  152. 171journalFormation of Amino Acids on the Sonolysis of Aqueous Solutions Containing Acetic Acid, Methane, or Carbon Dioxide, in the Presence of Nitrogen GasLeena Dharmarathne et al. — 7 January 2016
  153. 172journalCavitation as a plausible driving force for the prebiotic formation of N9 purine nucleosidesYeersen Patehebieke et al. — 2021
  154. 173journalCavitation-Induced Synthesis of Biogenic Molecules on Primordial EarthNatan-Haim Kalson et al. — 11 September 2017
  155. 174journalWere the first organisms heat engines? A new model for biogenesis and the early evolution of biological energy conversionAnthonie W. J. Muller — 1995
  156. 175journalThermal energy and the origin of lifeAnthonie W. J. Muller et al. — 2006
  157. 176journalATP SynthaseWolfgang Junge et al. — 2 June 2015
  158. 177bookThe Vital Question: Why Is Life The Way It Is?Nick Lane — Profile Books — 2015
  159. 178journalThe Hot Spring Hypothesis for an Origin of LifeBruce Damer et al. — 2020-04-01
  160. 180webWhy Comets Are Like Deep Fried Ice CreamWhitney Clavin — February 10, 2015
  161. 183journalWhen Did Life Likely Emerge on Earth in an RNA-First Process?S. A. Benner et al. — 2020
  162. 184journalThe 'Strong' RNA World Hypothesis: Fifty Years OldMarc Neveu et al. — 22 April 2013
  163. 185journalOrigin of life: The RNA worldWalter Gilbert — 20 February 1986
  164. 186journalThe origin of the RNA world: Co-evolution of genes and metabolismShelley D. Copley et al. — December 2007
  165. 187journalSome consequences of the RNA world hypothesisLeslie E. Orgel — April 2003
  166. 188harvnbRobertson, Joyce (2012)Robertson, Joyce — 2012
  167. 189harvnbNeveu, Kim, Benner (2013)Neveu, Kim, Benner — 2013
  168. 190journalThe origins of the RNA worldMichael P. Robertson et al. — May 2012
  169. 191journalThe RNA Worlds in ContextThomas R. Cech — July 2012
  170. 192journalOn the origin of life: an RNA-focused synthesis and narrativeJacob L. Fine et al. — August 2023
  171. 193journalGetting Past the RNA World: The Initial Darwinian AncestorMichael Yarus — April 2011
  172. 194harvnbVoet, Voet (2004) p. 29Voet, Voet — 2004
  173. 195journalOrigin and evolution of the ribosomeGeorge.E. Fox — 9 June 2010
  174. 196journalSelf-Sustained Replication of an RNA EnzymeTracey A. Lincoln et al. — 27 February 2009
  175. 197journalEvolution in an RNA worldGerald F. Joyce — 2009
  176. 198webThe Origins of Function in Biological Nucleic Acids, Proteins, and MembranesJack W. Szostak — Howard Hughes Medical Institute — 5 February 2015
  177. 199journalThe Darwinian DynamicHarris Bernstein et al. — June 1983
  178. 200harvnbMichod (1999)Michod — 1999
  179. 201arxivPrimordial RNA Replication and Applications in PCR TechnologyStan Palasek — 23 May 2013
  180. 202journalThe RNA World on Ice: A New Scenario for the Emergence of RNA InformationAlexander V. Vlassov et al. — August 2005
  181. 203journalThe emergence of the non-cellular phase of life on the fine-grained clayish particles of the early Earth's regolithMark D. Nussinov et al. — 1997
  182. 204journaltRNA sequences can assemble into a replicatorAlexandra Kühnlein et al. — 2 March 2021
  183. 206journalPhylogenetic structure of the prokaryotic domain: the primary kingdoms.C. R. Woese et al. — 1977
  184. 207journalThe origin of a derived superkingdom: how a gram-positive bacterium crossed the desert to become an archaeonR. E. Valas et al. — 2011
  185. 208journalRooting the tree of life by transition analysesThomas Cavalier-Smith — 2006
  186. 210journalEarly life liked it hot2016
  187. 211journalIs LUCA a thermophilic progenote?Johann Peter Gogarten et al. — 2016-11-25
  188. 214journalComparative genomics, minimal gene-sets and the last universal common ancestorE. V. Koonin — 2003
  189. 215journalA biophysical basis for the emergence of the genetic code in protocellsStuart A. Harrison et al. — 2022-11-01
  190. 216journalLife as a guide to prebiotic nucleotide synthesisStuart A. Harrison et al. — 2018-12-12
  191. 217journalEnvironmental Adaptation from the Origin of Life to the Last Universal Common AncestorMarjorie D. Cantine et al. — 2018-03-01
  192. 218journalThe genomics of LUCAWai-Kin Mat — May 1, 2008
  193. 219bookSecret Chambers: The Inside Story of Cells and Complex LifeM. D. Brasier — Oxford University Press — 2012
  194. 220journalHydrothermal vents and prebiotic chemistry: a reviewM. Colín-García et al. — 2016
  195. 221webHydrothermal Vents Could Explain Chemical Precursors to LifeMichael Schirber — NASA — 24 June 2014
  196. 222journalOn the origins of cells: a hypothesis for the evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from prokaryotes to nucleated cellsWilliam Martin et al. — 29 January 2003
  197. 223harvnbLane (2009)Lane — 2009
  198. 224press releaseChemistry of seabed's hot vents could explain emergence of lifeOli Usher — University College London — 27 April 2015
  199. 227journalSubmarine hydrothermal vents and associated gradient environments as sites for the origin and evolution of lifeJ. A. Baross et al. — 1985
  200. 228journalThe emergence of life from iron monosulphide bubbles at a submarine hydrothermal redox and pH frontM. J. Russell et al. — 1997
  201. 229journalThe energetics of organic synthesis inside and outside the cellJ. P. Amend et al. — 2013
  202. 230journalGeomicrobiology and microbial geochemistry:principles of geobiochemistryE. L. Shock et al. — 2015
  203. 231journalOn the origin of biochemistry at an alkaline hydrothermal ventW. Martin et al. — 2007
  204. 232journalRNA Oligomerization in Laboratory Analogues of Alkaline Hydrothermal Vent SystemsBradley T. Burcar et al. — 2015
  205. 233journalRNA Oligomerization in Laboratory Analogues of Alkaline Hydrothermal Vent SystemsBradley T. Burcar et al. — July 2015
  206. 234journalThe Origin of Membrane BioenergeticsNick Lane et al. — 2012-12-21
  207. 235journalExtreme accumulation of nucleotides in simulated hydrothermal pore systemsPhilipp Baaske et al. — 2007-05-29
  208. 236journalThe limits of metabolic heredity in protocellsRaquel Nunes Palmeira et al. — 2022-11-09
  209. 237journalBacterial evolutionCarl R. Woese — 1987
  210. 238bookEvolution of Early Earth's Atmosphere, Hydrosphere, and Biosphere – Constraints from Ore DepositsMichael J. Russell et al. — Geological Society of America — 2006
  211. 239journalParallel adaptations to high temperatures in the Archaean eonBastien Boussau et al. — December 2008
  212. 240journalA Nonhyperthermophilic Common Ancestor to Extant Life FormsNicolas Galtier et al. — 1999-01-08
  213. 241journalAstrophysical and astrochemical insights into the origin of lifeP. Ehrenfreund et al. — August 2002
  214. 242journalClumped isotopologue constraints on the origin of methane at seafloor hot springsDavid T. Wang et al. — February 2018
  215. 243bookComets and the Origin and Evolution of LifeC.F. Chyba et al. — Springer — 2006
  216. 244bookFrom Stardust to First Cells: The Origin and Evolution of Early LifeSankar Chatterjee — Springer International Publishing — 2023
  217. 245journalBiogeochemical explanations for the world's most phosphate-rich lake, an origin-of-life analogSebastian Haas et al. — 9 January 2024
  218. 246bookAssembling LifeDavid W. Deamer — Oxford University Press — 7 February 2019
  219. 247journalOrigin of the RNA world: The fate of nucleobases in warm little pondsBen K. D. Pearce et al. — 2 October 2017
  220. 248journalPromotion of protocell self-assembly from mixed amphiphiles at the origin of lifeSean F. Jordan et al. — 2019
  221. 249bookIs Earth Exceptional? The quest for cosmic lifeMario Livio et al. — Basic Books — 2024
  222. 250journalWas There Land on the Early Earth?Jun Korenaga — November 2021
  223. 251journalEmergence of a Habitable PlanetKevin Zahnle et al. — 2007-03-01
  224. 252journalOrigin of first cells at terrestrial, anoxic geothermal fieldsArmen Y. Mulkidjanian et al. — 2012-04-03
  225. 253journalSimple prebiotic synthesis of high diversity dynamic combinatorial polyester librariesKuhan Chandru et al. — 31 May 2018
  226. 254journalEster-Mediated Amide Bond Formation Driven by Wet–Dry Cycles: A Possible Path to Polypeptides on the Prebiotic EarthJay G. Forsythe et al. — 17 August 2015
  227. 255journalCommon origins of RNA, protein and lipid precursors in a cyanosulfidic protometabolismBhavesh H. Patel et al. — March 16, 2015
  228. 256journalWhere Did Life Begin? Testing Ideas in Prebiotic Analogue ConditionsDavid Deamer — 10 February 2021
  229. 257journalThe Archean atmosphereDavid C. Catling et al. — 2020-02-28
  230. 258journalThe origin of life?did it occur at high temperatures?Stanley L. Miller et al. — December 1995
  231. 259journalThe unique DNA topology and DNA topoisomerases of hyperthermophilic archaeaPatrick Forterre et al. — May 1996
  232. 260journalWidespread distribution of archaeal reverse gyrase in thermophilic bacteria suggests a complex history of vertical inheritance and lateral gene transfersCéline Brochier-Armanet et al. — May 2006
  233. 261journalThe faint young Sun problemGeorg Feulner — June 2012
  234. 262journalImpact melting of frozen oceans on the early Earth: Implications for the origin of lifeJ. L. Bada et al. — 1994-02-15
  235. 263journalInfluence of Ionic Inorganic Solutes on Self-Assembly and Polymerization Processes Related to Early Forms of Life: Implications for a Prebiotic Aqueous MediumPierre-Alain Monnard et al. — June 2002
  236. 264journalIn-ice evolution of RNA polymerase ribozyme activityJames Attwater et al. — December 2013
  237. 265journalRNA Folding Argues Against a Hot-Start Origin of LifeVincent Moulton et al. — 2000-10-01
  238. 266journalRNA folding in living cellsGeorgeta Zemora et al. — November 2010
  239. 267journalHypothesis: Origin of Life in Deep-Reaching Tectonic FaultsUlrich Schreiber et al. — 2012
  240. 268journalOrganic compounds in fluid inclusions of Archean quartz—Analogues of prebiotic chemistry on early EarthUlrich Schreiber et al. — 2017-06-14
  241. 269journalAliphatic Aldehydes in the Earth's Crust—Remains of Prebiotic Chemistry?Yildiz Großmann et al. — 2022-06-21
  242. 270journalSelection of Prebiotic Molecules in Amphiphilic EnvironmentsChristian Mayer et al. — 2017-01-07
  243. 271journalEmergence of homochirality in far-from-equilibrium systems: Mechanisms and role in prebiotic chemistryRaphaël Plasson et al. — August 2007
  244. 272harvnbChaichian, Rojas, Tureanu (2014) p. 353–364Chaichian, Rojas, Tureanu — 2014
  245. 274journalNoise-induced mechanism for biological homochirality of early life self-replicatorsFarshid Jafarpour et al. — 2015
  246. 275journalOn spontaneous asymmetric synthesisF. C. Frank — 1953
  247. 276journalPolarized Starlight and the Handedness of LifeStuart Clark — July–August 1999
  248. 277journalExtended High Circular Polarization in the Orion Massive Star Forming Region: Implications for the Origin of Homochirality in the Solar SystemTsubasa Fukue et al. — June 2010
  249. 278journalMolecular Asymmetry in Prebiotic Chemistry: An Account from MeteoritesSandra Pizzarello — 13 April 2016
  250. 279journalHighly Enantioselective Catalytic Asymmetric Automultiplication of Chiral Pyrimidyl AlcoholTakanori Shibata et al. — 17 January 1996
  251. 280journalAsymmetric autocatalysis and the origin of chiral homogeneity in organic compoundsKenso Soai et al. — 2001