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Abiogenesis

In biology, abiogenesis, or informally the origin


of life (OoL),[3][4][5][a] is the natural process by
which life has arisen from non-living matter, such
as simple organic compounds.[6][4][7][8] While
the details of this process are still unknown, the
prevailing scientific hypothesis is that the
transition from non-living to living entities was
not a single event, but an evolutionary process of
increasing complexity that involved molecular
self-replication, self-assembly, autocatalysis, and
the emergence of cell membranes.[9][10][11]
Although the occurrence of abiogenesis is
uncontroversial among scientists, its possible
mechanisms are poorly understood. There are
The earliest known life-forms are putative fossilized
several principles and hypotheses for how
microorganisms, found in hydrothermal vent
abiogenesis could have occurred.[12]
precipitates, that may have lived as early as 4.28 Gya
(billion years ago), relatively soon after the oceans
The study of abiogenesis aims to determine how
formed 4.41 Gya, and not long after the formation of the
pre-life chemical reactions gave rise to life under
Earth 4.54 Gya.[1][2]
conditions strikingly different from those on
Earth today.[13] It primarily uses tools from
biology, chemistry, and geophysics,[14] with
more recent approaches attempting a synthesis of all three:[15] more specifically, astrobiology, biochemistry,
biophysics, geochemistry, molecular biology, oceanography and paleontology. Life functions through the
specialized chemistry of carbon and water and builds largely upon four key families of chemicals: lipids
(cell membranes), carbohydrates (sugars, cellulose), amino acids (protein metabolism), and nucleic acids
(DNA and RNA). Any successful theory of abiogenesis must explain the origins and interactions of these
classes of molecules.[16] Many approaches to abiogenesis investigate how self-replicating molecules, or
their components, came into existence. Researchers generally think that current life descends from an RNA
world,[17] although other self-replicating molecules may have preceded RNA.[18][19]

The classic 1952 Miller–Urey experiment and similar research demonstrated that most amino acids, the
chemical constituents of the proteins used in all living organisms, can be synthesized from inorganic
compounds under conditions intended to replicate those of the early Earth. Scientists have proposed various
external sources of energy that may have triggered these reactions, including lightning and radiation. Other
approaches ("metabolism-first" hypotheses) focus on understanding how catalysis in chemical systems on
the early Earth might have provided the precursor molecules necessary for self-replication.[20]

The alternative panspermia hypothesis[21] speculates that microscopic life arose outside Earth by unknown
mechanisms, and spread to the early Earth on space dust[22] and meteoroids.[23] It is known that complex
organic molecules occur in the Solar System and in interstellar space, and these molecules may have
provided starting material for the development of life on Earth.[24][25][26][27] It has also been suggested that
life arises in a Great Prebiotic Spot on a given world.[28]
Earth remains the only place in the universe known to
harbour life,[29][30] and fossil evidence from the Earth
informs most studies of abiogenesis. The age of the Earth is
4.54 Gy (billion year);[31][32][33] the earliest undisputed
evidence of life on Earth dates from at least 3.5 Gya (Gy
ago),[34][35][36] and possibly as early as the Eoarchean Era
(3.6–4.0 Gya). In 2017 scientists found possible evidence of
early life on land in 3.48 Gyo (Gy old) geyserite and other
related mineral deposits (often found around hot springs and
geysers) uncovered in the Pilbara Craton of Western
Australia.[37][38][39][40] However, a number of discoveries
suggest that life may have appeared on Earth even earlier.
As of 2017, microfossils (fossilised microorganisms) within
hydrothermal-vent precipitates dated 3.77 to 4.28 Gya in Miller–Urey experiment Synthesis of small
rocks in Quebec may harbour the oldest record of life on organic molecules in a mixture of simple
Earth, suggesting life started soon after ocean formation 4.4 gases that is placed in a thermal gradient
Gya during the Hadean Eon.[1][2][41][42][43] by heating (right) and cooling (left) the
mixture at the same time, a mixture that is
The NASA strategy on abiogenesis states that it is necessary also subject to electrical discharges
to identify interactions, intermediary structures and
functions, energy sources, and environmental factors that
contributed to the diversity, selection, and replication of evolvable macromolecular systems.[44] Emphasis
must continue to map the chemical landscape of potential primordial informational polymers. The advent of
polymers that could replicate, store genetic information, and exhibit properties subject to selection likely
was a critical step in the emergence of prebiotic chemical evolution.[44]

Contents
Current life, the result of abiogenesis: biology
Definition of life
Fermentation
Chemiosmosis
ATP synthase
RNA world
Phylogeny and LUCA
Key issues in abiogenesis
Protein vs. nucleic acid as the precursor to protein synthesis
Emergence of the genetic code
Error in translation catastrophe
Homochirality
Early universe and Earth
Early universe with first stars
Emergence of Earth
Earliest evidence of life: palaeontology
Conceptual history until the 1960s: biology
Panspermia
Origin of life posited directly after the Big Bang and have spread over the entire
Universe
Panspermia by life brought from Mars to Earth
Spontaneous generation
General acceptance of spontaneous generation until the 19th century
Spontaneous generation considered disproven in the 19th century
Etymology of biogenesis and abiogenesis
Oparin: Primordial soup hypothesis
John Bernal
Miller–Urey experiment
Primordial origin of biological molecules: Chemistry
Observed extraterrestrial organic molecules
Amino acids
PAH world hypothesis
Nucleobases
The sugar glycolaldehyde
Polyphosphates
Chemical synthesis in the laboratory
Fox proteinoids
Sugars
Nucleobases
Use of reactions for ammonium cyanide
Effects with temperatures around the freezing point of water
Use of less-reducing gas in Miller–Urey experiment
Synthesis based on hydrogen cyanide
Issues during laboratory synthesis
Autocatalysis
Pertinent geological environments
Darwin's little pond
Volcanic hot springs and hydrothermal vents, shallow or deep
Deep sea hydrothermal vents
Fluctuating hydrothermal pools on volcanic islands or proto-continents
Volcanic ash in the ocean
Gold's deep-hot biosphere
Radioactive beach hypothesis
Thermodynamics, self-organization, and information: Physics
Thermodynamics principles: Energy and entropy
Obtaining free energy
Self-organization
Dissipative structuring
Self-organization by dissipative structures
Encapsulation: morphology
Encapsulation without a membrane
Oparin's coacervate
Membraneless polyester droplets
Proteinoid microspheres
Lipid world
Protocells
Lipid vesicles formation in fresh water
Vesicles consisting of mixtures of RNA-like biochemicals
Metal-sulfide precipitates
Origin of metabolism: physiology
Clay hypothesis
Iron–sulfur world
Zinc-world hypothesis
Other abiogenesis scenarios
Chemical pathways described by computer
The hypercycle
Organic pigments in dissipative structures
Protein amyloid
Fluctuating salinity: dilute and dry-down
A first protein that condenses substrates during thermal cycling: thermosynthesis
Emergence of chemiosmotic machinery
Pre-RNA world: The ribose issue and its bypass
RNA structures
Viral origin
RNA world
RNA-DNA world
Experiments on the origin of life
See also
References
Sources
Further reading
External links

Current life, the result of abiogenesis: biology

Definition of life

As many as 123 definitions of life have been compiled.[45]

The definition of life is somewhat disagreed upon; different biology textbooks define life differently. James
Gould notes:

Most dictionaries define life as the property that distinguishes the living from the dead, and
define dead as being deprived of life. These singularly circular and unsatisfactory definitions
give us no clue to what we have in common with protozoans and plants.[46]
Neil Campbell and Jane Reece wrote:

The phenomenon we call life defies a simple, one-sentence definition.[47]

This difference can also be found in books on the origin of life. John Casti gives a single-sentence
definition:

By more or general consensus nowadays, an entity is considered to be "alive" if it has the


capacity to carry out three basic functional activities: metabolism, self-repair, and replication.
[48]

In contrast, Dirk Schulze-Makuch and Louis Irwin devote the entire first chapter of their book to discussing
a definition of life.[49]

Nonetheless, a definition of life currently favored by NASA is that life is "a self-sustaining chemical system
capable of Darwinian evolution."[50][51][52][53] More simply, life is "matter that can reproduce itself and
evolve as survival dictates".[54][55][56]

Fermentation

Citric acid cycle

Albert Lehninger has stated around 1970 that fermentation,


including glycolysis, is a suitable primitive energy source for the
origin of life.[57]

Since living organisms probably first arose in an


atmosphere lacking oxygen, anaerobic fermentation is Overall diagram of the chemical
the simplest and most primitive type of biological reactions of metabolism, in which the
mechanism for obtaining energy from nutrient citric acid cycle can be recognized
molecules. as the circle just below the middle of
the figure

Fermentation involves glycolysis, which transduces the chemical


energy of sugar into the chemical energy of ATP.

Chemiosmosis

As fermentation had been elucidated around 1970, whereas the mechanism of oxidative phosphorylation
had not, and some controversies still existed, processes other than fermentation may have looked too
complex at that time. Peter Mitchell's chemiosmosis is now however generally accepted as correct.
Even Peter Mitchell himself assumed that fermentation
preceded chemiosmosis. Chemiosmosis is, however,
ubiquitous in life. A model for the origin of life has been
presented in terms of chemiosmosis. [58][59]

Both respiration by mitochondria and photosynthesis in


chloroplasts make use of chemiosmosis to generate most
of their ATP.

Oxidative phosphorylation Today the energy sources of almost all life can be linked
to photosynthesis, and one speaks of primary production
by sunlight. The oxygen that powers organisms[60] that
oxidize H2 or H2 S at hydrothermal vents at the bottom of
the ocean is the result of photosynthesis at the oceans'
surface.

ATP synthase

The mechanism of ATP synthesis is complex and


involves a closed membrane in which the ATP synthase
is embedded. The ATP is synthesized by the F1 subunit
Chemiosmotic coupling mitochondrion of ATP synthase by the binding change mechanism
discovered by Paul Boyer. The energy required to release
formed strongly-bound
ATP has its origin in protons that move
across the membrane. These protons have
been set across the membrane during
respiration or photosynthesis.

RNA world

The RNA world hypothesis describes an


early Earth with self-replicating and
catalytic RNA but no DNA or
proteins.[62] It is widely accepted that
current life on Earth descends from an
RNA world,[17][63][64] although RNA- Paul Boyer
based life may not have been the first life
to exist.[18][19] This conclusion is drawn
from many independent lines of evidence, such as the observations that
Depiction of ATP synthase
RNA is central to the translation process and that small RNAs can catalyze
using the chemiosmotic
proton gradient to power
all of the chemical groups and information transfers required for
ATP synthesis through life.[19][65] The structure of the ribosome has been called the "smoking
oxidative phosphorylation. gun," as it showed that the ribosome is a ribozyme, with a central core of
RNA and no amino acid side chains within 18 angstroms of the active site
where peptide bond formation is catalyzed.[18][66] Nonetheless, in March
2021, researchers reported evidence suggesting that a preliminary form of transfer RNA could have been a
replicator molecule itself in the very early development of life.[67][68]
The concept of the RNA world was first proposed in
1962 by Alexander Rich,[69] and the term was coined by
Walter Gilbert in 1986.[19][70] In March 2020,
astronomer Tomonori Totani presented a statistical
approach for explaining how an initial active RNA
molecule might have been produced randomly in the
universe sometime since the Big Bang.[71][72]

Phylogeny and LUCA

Starting with the work of Carl Woese, molecular studies


have placed the last universal common ancestor (LUCA)
between Bacteria and a clade formed by Archaea and
Eukaryota in the phylogenetic tree of life.[73][74] A
minority of studies have placed the LUCA in Bacteria,
Molecular structure of the ribosome 30S
proposing that archaea and eukaryotes are evolutionarily
subunit from Thermus thermophilus.[61]
derived from within eubacteria.[75] Thomas Cavalier- Proteins are shown in blue and the single RNA
Smith hypothesizes that the phenotypically diverse chain in orange.
phylum Chloroflexi contained the LUCA.[76]

In 2005, Peter Ward proposed that


abiotically synthesized RNA became
enclosed within a capsule and then created
RNA ribozyme replicates. This then
bifurcated between Dominion Ribosa (RNA
life), Domain Viorea (Viruses), and
Dominion Terroa (Cellular life), which
contains the LUCA of earlier phylogenic
trees.[77]

In 2016, a set of 355 genes likely present in


the Last Universal Common Ancestor
(LUCA) of all organisms living on Earth
was identified.[78] A total of 6.1 million
A cladogram placing other extreme hyperthermophiles
prokaryotic genes from Bacteria and Archae
alongside the LUCA at the base of the phylogenetic tree of
were sequenced, identifying 355 protein
life.
clusters from amongst 286,514 protein
clusters that were probably common to
LUCA. The result suggest that the LUCA
was anaerobic with a Wood–Ljungdahl pathway, nitrogen- and carbon-fixing, thermophilic. Its cofactors
suggest dependence upon an environment rich in hydrogen, carbon dioxide, iron, and transition metals. Its
genetic code required nucleoside modifications and methylation. LUCA likely inhabited an anaerobic
hydrothermal vent setting in a geochemically active environment.[79][80]

Key issues in abiogenesis

Protein vs. nucleic acid as the precursor to protein synthesis


Possible precursors for the evolution of protein synthesis include a mechanism to synthesize short peptide
cofactors or form a mechanism for the duplication of RNA. It is likely that the ancestral ribosome was
composed entirely of RNA, although some roles have since been taken over by proteins. Major remaining
questions on this topic include identifying the selective force for the evolution of the ribosome and
determining how the genetic code arose.[81]

Eugene Koonin said,

Despite considerable experimental and theoretical effort, no compelling scenarios currently


exist for the origin of replication and translation, the key processes that together comprise the
core of biological systems and the apparent pre-requisite of biological evolution. The RNA
World concept might offer the best chance for the resolution of this conundrum but so far
cannot adequately account for the emergence of an efficient RNA replicase or the translation
system. The MWO ["many worlds in one"] version of the cosmological model of eternal
inflation could suggest a way out of this conundrum because, in an infinite multiverse with a
finite number of distinct macroscopic histories (each repeated an infinite number of times),
emergence of even highly complex systems by chance is not just possible but inevitable.[82]

Emergence of the genetic code

See: Genetic code.

Error in translation catastrophe

Hoffmann has shown that an early error-prone translation machinery can be stable against an error
catastrophe of the type that had been envisaged as problematical for the origin of life, and was known as
"Orgel's paradox".[83][84][85]

Homochirality

Homochirality refers to a geometric uniformity of some materials composed of chiral units. Chiral refers to
nonsuperimposable 3D forms that are mirror images of one another, as are left and right hands. Living
organisms use molecules that have the same chirality ("handedness"): with almost no exceptions,[86] amino
acids are left-handed while nucleotides and sugars are right-handed. Chiral molecules can be synthesized,
but in the absence of a chiral source or a chiral catalyst, they are formed in a 50/50 mixture of both
enantiomers (called a racemic mixture). Known mechanisms for the production of non-racemic mixtures
from racemic starting materials include: asymmetric physical laws, such as the electroweak interaction;
asymmetric environments, such as those caused by circularly polarized light, quartz crystals, or the Earth's
rotation, statistical fluctuations during racemic synthesis,[87] and spontaneous symmetry
breaking.[88][89][90]

Once established, chirality would be selected for.[91] A small bias (enantiomeric excess) in the population
can be amplified into a large one by asymmetric autocatalysis, such as in the Soai reaction.[92] In
asymmetric autocatalysis, the catalyst is a chiral molecule, which means that a chiral molecule is catalyzing
its own production. An initial enantiomeric excess, such as can be produced by polarized light, then allows
the more abundant enantiomer to outcompete the other.[93]
Clark has suggested that homochirality may have started in outer space, as the studies of the amino acids on
the Murchison meteorite showed that L-alanine is more than twice as frequent as its D form, and L-
glutamic acid was more than three times prevalent than its D counterpart. Various chiral crystal surfaces can
also act as sites for possible concentration and assembly of chiral monomer units into
macromolecules.[94][95] Compounds found on meteorites suggest that the chirality of life derives from
abiogenic synthesis, since amino acids from meteorites show a left-handed bias, whereas sugars show a
predominantly right-handed bias, the same as found in living organisms.[96]

Early universe and Earth

Early universe Nature timeline


with first stars 0— ← Earliest apes / humans
Vertebrates ← Earliest mammals
– ← Cambrian explosion
Soon after the Big Bang, ← Earliest animals / plants
which occurred roughly −1 — Multicellular
14 Gya, the only chemical life
– L ← Earliest fungi
elements present in the i
universe were hydrogen, −2 — f ← Earliest multicellular life
e
helium, and lithium, the – ← Atmospheric oxygen
three lightest atoms in the Photosynthesis
periodic table. These −3 —
elements gradually came – ← Earliest oxygen
together to form stars. Single-celled life
These early stars were −4 — ← Earliest life
Water
massive and short-lived,
– ← Earth / Solar System
producing heavier elements
through stellar −5 — Accelerated expansion
nucleosynthesis. Carbon,

currently the fourth most
abundant chemical element −6 —
in the universe (after
hydrogen, helium and
– ← Alpha Centauri
← NGC 188 star cluster
oxygen), was formed −7 —
mainly in white dwarf
stars, particularly those –
bigger than two solar −8 — Matter-dominated
masses.[97][98] era

As these stars reached the ← Milky Way spirals
−9 —
end of their lifecycles, they
ejected these heavier –
elements, among them −10 — ← Andromeda Galaxy
carbon and oxygen,
throughout the universe. –
These heavier elements −11 —
allowed for the formation
of new objects, including – ← Omega Centauri
rocky planets and other
−12 —
bodies.[99]

3
According to the nebular −13 — ← Quasar / black hole
Reionization
hypothesis, the formation ← Earliest galaxy
and evolution of the Solar Dark Ages ← Earliest stars
← Earliest Universe
System began 4.6 Gya (billion years ago)
with the gravitational
collapse of a small part of a
giant molecular cloud.[100] Most of the collapsing mass collected in the center, forming the Sun, while the
rest flattened into a protoplanetary disk out of which the planets, moons, asteroids, and other small Solar
System bodies formed.

Life timeline
0— ← Quaternary ice age*
Emergence of P Flowers Birds Primates ← Earliest apes / humans
–h Mammals
Earth a Dinosaurs
—n
e P ← Karoo ice age*
The Hadean Earth was at –r l Arthropods Molluscs ← Earliest tetrapods
first inhospitable to any o a ← Andean glaciation*
−500 — z n ← Cambrian explosion
living organisms. During – oi t Ediacaran biota
s ←
← Cryogenian ice age*
its formation, the Earth lost —c ← Earliest animals
a significant part of its ← Earliest plants

initial mass, and
consequentially lacked the −1000 —
Multicellular life
gravity to hold molecular –
hydrogen and the bulk of —
the original inert P
–r
gases.[101] The atmosphere −1500 — ot ← Earliest fungi
consisted largely of water ← Earliest multicellular life
– er
vapor, nitrogen and carbon
—o
dioxide, with smaller z
–o Eukaryotes
amounts of carbon i
monoxide, hydrogen, and −2000 — c
sulfur compounds.[102] –
The solution of carbon — ← Huronian glaciation*
dioxide in water is thought ← Atmospheric oxygen

to have made the seas
−2500 —
slightly acidic, giving them
a pH of about 5.5.[103] The –
Hadean atmosphere has —
Photosynthesis ← Pongola glaciation*
been characterized as a –
"gigantic, productive −3000 —
outdoor chemical A
–r
laboratory," [104] similar to c
—h
volcanic gases today which
–ea
still support some abiotic
chemistry.[104] −3500 — n ← Earliest oxygen

Oceans may have appeared — Single-celled life
as soon as 200 My after the – ← LHB meteorites
Earth formed, in a hot,
−4000 — ← Earliest life
100 C, reducing H
–a
environment, as the pH of d Water
—e
e
5.8 rose rapidly towards –a
neutral.[105] This scenario n ← Earliest water
has found support from the
−4500 — ← Earth formed
(4540 mya)
dating of 4.404 Gya zircon (million years ago) *Ice Ages
crystals from
metamorphosed quartzite of Mount Narryer in Western Australia, which provide evidence that oceans and
continental crust existed within 150 Ma of Earth's formation.[106] Despite the likely increased volcanism
and existence of many smaller tectonic "platelets," it has been suggested that between 4.4 and 4.3 Gya, the
Earth was a water world, with little if any continental crust, an extremely turbulent atmosphere and a
hydrosphere subject to intense ultraviolet (UV) light, from a T Tauri stage Sun, cosmic radiation and
continued bolide impacts.[107] Internal heating as a result of gravitational sorting between the core and the
mantle would have caused a great deal of mantle convection, with the probable result of many more smaller
and more active tectonic plates than now exist.

The hypothesis for the Late Heavy Bombardment posits that the Hadean environment between 4.28[1][2]
and 3.8 Gya would have been highly hazardous to modern life. Following the Nice model, changes in the
orbits of the giant planets may have bombarded the Earth with asteroids and comets that pockmarked the
Moon and inner planets.[108] Frequent stellar collisions with objects up to 500 km in diameter would
sterilize the planet's surface and vaporize the oceans within a few months of impact. Hot steam and rock
vapor formed high altitude clouds that would completely cover the planet,[104] making photosynthesis
unviable. Rains would slowly have drawn down the height of the clouds, returning the oceans to their
original depth only 3,000 y after the impact event.[109] Impacts before 3.5 Gya would have also brought
quantities of organics comparable to those produced by terrestrial sources.[110][111] The periods between
such devastating environmental events give time windows for the possible origin of life in early
environments. If the deep marine hydrothermal setting was the site for the origin of life, then abiogenesis
could have happened as early as 4.0-4.2 Gya. If the site was at the surface of the Earth, abiogenesis could
only have occurred between 3.7 and 4.0 Gya.[112] However, new lunar surveys and samples have led
scientists, including an architect of the Nice model, to deemphasize the LHB.[113]

If life evolved deeper than ten meters, it would have been shielded from both late impacts and high levels of
UV radiation from the T Tauri stage Sun. Simulations of geothermically heated oceanic crust yield far more
organics than those found in the Miller–Urey experiments. In the deep hydrothermal vents, Everett Shock
has found "there is an enormous thermodynamic drive to form organic compounds, as seawater and
hydrothermal fluids, which are far from equilibrium, mix and move towards a more stable state."[114]
Shock found that the available energy is maximized at 100–150 °C, precisely the temperatures at which the
hyperthermophilic bacteria and thermoacidophilic archaea have been found living. These organisms are
placed at the base of the phylogenetic tree of life, closest to the Last Universal Common Ancestor
(LUCA).[115]

Earliest evidence of life: palaeontology


The earliest life on Earth existed more than 3.5 Gya (billion years ago),[34][35][36] during the Eoarchean
Era when sufficient crust had solidified following the molten Hadean Eon. The earliest physical evidence
so far found consists of microfossils in the Nuvvuagittuq Greenstone Belt of Northern Quebec, in banded
iron formation rocks at least 3.77 and possibly 4.28 Gya.[1][116] This finding suggested life developed very
soon after oceans formed. The structure of the microbes was noted to be similar to bacteria found near
hydrothermal vents in the modern era, and provided support for the hypothesis that abiogenesis began near
hydrothermal vents.[42][1]
Biogenic graphite has
been found in 3.7 Gyo
metasedimentary rocks
from southwestern
Greenland [117] and
microbial mat fossils
found in 3.48 Gyo
sandstone from Western
Australia.[118][119]
Precambrian stromatolites in the
Siyeh Formation, Glacier National
Evidence of early life in Stromatolites in Shark Bay
Park. A 2002 study suggested that rocks from Akilia Island,
these 3.5 Gya (3.5 billion year old) near the Isua supracrustal
formations. This suggests they are belt in southwestern Greenland, dating to 3.7 Gya have shown
evidence of one of the earliest life biogenic carbon isotopes.[120][121] In other parts of the Isua
forms on Earth. supracrustal belt, graphite inclusions trapped within garnet crystals
are connected to the other elements of life: oxygen, nitrogen, and
possibly phosphorus in the form of phosphate, providing further
evidence for life 3.7 Gya. [122] At Strelley Pool, in the Pilbara region of Western Australia, compelling
evidence of early life was found in pyrite-bearing sandstone in a fossilized beach, that showed rounded
tubular cells that oxidized sulfur by photosynthesis in the absence of oxygen.[123][124][125] Further research
on zircons from Western Australia in 2015 suggested that life likely existed on Earth at least 4.1
Gya.[126][127][128]

In 2019 Raphael Baumgartner at the University of New South Wales in Australia and his colleagues looked
at rocks in the Pilbara region of Western Australia. This area contains some of the oldest preserved rocks on
Earth. Of the three most important sites, the Dresser Formation is the oldest, with rocks that are 3.48 billion
years old. The Dresser Formation appears to contain layered structures called stromatolites.[129] These
stromatolites lie within undeformed hydrothermal-sedimentary strata and show textural features that are
indicative of biogenic origins. In 2017, Tara Djokic and her team showed that parts of the Dresser
formation preserve hot spring on lands, but other regions seem to have been shallow seas.[130]

Conceptual history until the 1960s: biology

Panspermia

Panspermia is the hypothesis that life exists throughout the universe, distributed by meteoroids, asteroids,
comets[131] and planetoids.[132]

The panspermia hypothesis does not attempt to explain how life first originated but merely shifts the origin
to another planet or a comet. The advantage of an extraterrestrial origin of primitive life is that life is not
required to have formed on each planet it occurs on, but rather in a single location, and then spread about
the galaxy to other star systems via cometary or meteorite impact.[133] Evidence for the panspermia
hypothesis is scant, but it finds some support in studies of Martian meteorites found in Antarctica and in
studies of extremophile microbes' survival in outer space tests.[134][135][136][137]

In August 2020, scientists reported that bacteria from Earth, particularly Deinococcus radiodurans, which
is highly resistant to environmental hazards, were found to survive for three years in outer space, based on
studies conducted on the International Space Station.[138][139]

Origin of life posited directly after the Big Bang and have spread over the entire Universe
An extreme speculation is that the biochemistry of life could have begun as early as 17 My (million years)
after the Big Bang, during a habitable epoch, and that life may exist throughout the universe.[140][141]

Panspermia by life brought from Mars to Earth

Carl Zimmer has speculated that the chemical conditions, including the presence of boron, molybdenum
and oxygen needed for the initial production of RNA, may have been better on early Mars than on early
Earth.[142][143][144] If so, life-suitable molecules originating on Mars may have later migrated to Earth via
meteor ejections.

Spontaneous generation

General acceptance of spontaneous generation until the 19th century

Traditional religion attributed the origin of life to supernatural deities who created the natural world.
Spontaneous generation, the first naturalistic theory of life arising from non-life, goes back to Aristotle and
ancient Greek philosophy, and continued to have support in Western scholarship until the 19th century.[145]
Classical notions of spontaneous generation held that certain "lower" or "vermin" animals are generated by
decaying organic substances. According to Aristotle, it was readily observable that aphids arise from dew
on plants, flies from putrid matter, mice from dirty hay, crocodiles from rotting sunken logs, and so on.[146]
A related theory was heterogenesis: that some forms of life could arise from different forms (e.g. bees from
flowers).[147] The modern scientist John Bernal said that the basic idea of such theories was that life was
continuously created as a result of chance events.[148]

In the 17th century, people began to question such assumptions. In 1646, Thomas Browne published his
Pseudodoxia Epidemica (subtitled Enquiries into Very many Received Tenets, and commonly Presumed
Truths), which was an attack on false beliefs and "vulgar errors." His contemporary, Alexander Ross,
erroneously refuted him, stating:

To question this [spontaneous generation], is to question Reason, Sense, and Experience: If he


doubts of this, let him go to Ægypt, and there he will find the fields swarming with mice begot
of the mud of Nylus, to the great calamity of the Inhabitants.[149][150]

In 1665, Robert Hooke published the first drawings of a microorganism.


Hooke was followed in 1676 by Antonie van Leeuwenhoek, who drew
and described microorganisms that are now thought to have been protozoa
and bacteria.[151] Many felt the existence of microorganisms was evidence
in support of spontaneous generation, since microorganisms seemed too
simplistic for sexual reproduction, and asexual reproduction through cell
division had not yet been observed. Van Leeuwenhoek took issue with the
ideas common at the time that fleas and lice could spontaneously result
from putrefaction, and that frogs could likewise arise from slime. Using a
broad range of experiments ranging from sealed and open meat incubation
and the close study of insect reproduction he became, by the 1680s,
convinced that spontaneous generation was incorrect.[152] Antonie van Leeuwenhoek

The first experimental evidence against spontaneous generation came in


1668 when Francesco Redi showed that no maggots appeared in meat when flies were prevented from
laying eggs. It was gradually shown that, at least in the case of all the higher and readily visible organisms,
the previous sentiment regarding spontaneous generation was false. The alternative hypothesis was
biogenesis: that every living thing came from a pre-existing living thing (omne vivum ex ovo, Latin for
"every living thing from an egg").[153] In 1768, Lazzaro Spallanzani demonstrated that microbes were
present in the air, and could be killed by boiling. In 1861, Louis Pasteur performed a series of experiments
that demonstrated that organisms such as bacteria and fungi do not spontaneously appear in sterile, nutrient-
rich media, but could only appear by invasion from without.

Spontaneous generation considered disproven in the 19th century

By the middle of the 19th century,


biogenesis had accumulated so much
evidence in support that the alternative
theory of spontaneous generation had
been effectively disproven. Pasteur
remarked, about a finding of his in 1864
which he considered definitive,

Never will the doctrine of


spontaneous generation
recover from the mortal blow
struck by this simple Charles Darwin in 1879
Louis Pasteur
experiment.[154][155]

gave a mechanism by which life diversified from a few simple organisms to a variety of to complex forms.
Today, scientists agree that all current life descends from earlier life, which has become progressively more
complex and diverse through Charles Darwin's mechanism of evolution by natural selection. Darwin wrote
to J.D. Hooker on 29 March 1863 stating that,

It is mere rubbish, thinking at present of the origin of life; one might as well think of the origin
of matter.

In On the Origin of Species, he had referred to life having been "created", by which he "really meant
'appeared' by some wholly unknown process", but had soon regretted using the Old Testament term
"creation".[156]

Etymology of biogenesis and abiogenesis

The term biogenesis is usually credited to either Henry Bastian or to Thomas Huxley.[157] Bastian used the
term around 1869 in an unpublished exchange with John Tyndall to mean "life-origination or
commencement". In 1870, Huxley, as new president of the British Association for the Advancement of
Science, delivered an address entitled Biogenesis and Abiogenesis.[158] In it he introduced the term
biogenesis (with an opposite meaning to Bastian's) as well as abiogenesis:

And thus the hypothesis that living matter always arises by the agency of pre-existing
living matter, took definite shape; and had, henceforward, a right to be considered and a
claim to be refuted, in each particular case, before the production of living matter in any
other way could be admitted by careful reasoners. It will be necessary for me to refer to this
hypothesis so frequently, that, to save circumlocution, I shall call it the hypothesis of
Biogenesis; and I shall term the contrary doctrine—that living matter may be produced by
not living matter—the hypothesis of Abiogenesis.[158]

Subsequently, in the preface to Bastian's 1871 book, The Modes of Origin of Lowest Organisms,[159]
Bastian referred to the possible confusion with Huxley's usage and explicitly renounced his own meaning:

A word of explanation seems necessary with regard to the introduction of the new term
Archebiosis. I had originally, in unpublished writings, adopted the word Biogenesis to
express the same meaning—viz., life-origination or commencement. But in the meantime,
the word Biogenesis has been made use of, quite independently, by a distinguished
biologist [Huxley], who wished to make it bear a totally different meaning. He also
introduced the word Abiogenesis. I have been informed, however, on the best authority,
that neither of these words can—with any regard to the language from which they are
derived—be supposed to bear the meanings which have of late been publicly assigned to
them. Wishing to avoid all needless confusion, I therefore renounced the use of the word
Biogenesis, and being, for the reason just given, unable to adopt the other term, I was
compelled to introduce a new word, in order to designate the process by which living
matter is supposed to come into being, independently of pre-existing living matter.[160]

Since the end of the nineteenth century, 'evolutive abiogenesis' means increasing complexity and evolution
of matter from inert to living states.[161]

Oparin: Primordial soup hypothesis

There is no single generally accepted model for the origin of life. Scientists have proposed several plausible
hypotheses which share some common elements. While differing in details, these hypotheses are based on
the framework laid out by Alexander Oparin (in 1924) and John Haldane (in 1925), that the first molecules
constituting the earliest cells

. . . were synthesized under natural conditions by a slow process of molecular evolution, and
these molecules then organized into the first molecular system with properties with biological
order".[162]

Oparin and Haldane suggested that the atmosphere of the early Earth may have been chemically reducing
in nature, composed primarily of methane (CH4 ), ammonia (NH3 ), water (H2 O), hydrogen sulfide (H2 S),
carbon dioxide (CO2 ) or carbon monoxide (CO), and phosphate (PO4 3−), with molecular oxygen (O2 ) and
ozone (O3 ) either rare or absent. According to later models, the atmosphere in the late Hadean period
consisted largely of nitrogen (N2 ) and carbon dioxide, with smaller amounts of carbon monoxide, hydrogen
(H2 ), and sulfur compounds;[163] while it did lack molecular oxygen and ozone,[164] it was not as
chemically reducing as Oparin and Haldane supposed.

No new notable research or hypothesis on the subject appeared until 1924, when Oparin reasoned that
atmospheric oxygen prevents the synthesis of certain organic compounds that are necessary building blocks
for life. In his book The Origin of Life,[165][166] he proposed (echoing Darwin) that the "spontaneous
generation of life" that had been attacked by Pasteur did, in fact, occur once, but was now impossible
because the conditions found on the early Earth had changed, and preexisting organisms would
immediately consume any spontaneously generated organism. Oparin argued that a "primeval soup" of
organic molecules could be created in an oxygenless atmosphere through the action of sunlight. These
would combine in ever more complex ways until they formed coacervate droplets. These droplets would
"grow" by fusion with other droplets, and "reproduce" through fission into daughter droplets, and so have a
primitive metabolism in which factors that promote "cell integrity" survive, and those that do not become
extinct. Many modern theories of the origin of life still take Oparin's ideas as a starting point.

About this time, Haldane suggested that the Earth's prebiotic oceans (quite different from their modern
counterparts) would have formed a "hot dilute soup" in which organic compounds could have formed.
Bernal called this idea biopoiesis or biopoesis, the process of living matter evolving from self-replicating
but non-living molecules,[148][167] and proposed that biopoiesis passes through a number of intermediate
stages.

Robert Shapiro has summarized the "primordial soup" theory of Oparin and Haldane in its "mature form"
as follows:[168]

1. The early Earth had a chemically reducing atmosphere.


2. This atmosphere, exposed to energy in various forms, produced simple organic compounds
("monomers").
3. These compounds accumulated in a "soup" that may have concentrated at various locations
(shorelines, oceanic vents etc.).
4. By further transformation, more complex organic polymers—and ultimately life—developed
in the soup.

John Bernal

John Bernal showed that based upon this and subsequent work there is no difficulty in principle in forming
most of the molecules we recognize as the necessary molecules for life from their inorganic precursors. The
underlying hypothesis held by Oparin, Haldane, Bernal, Miller and Urey, for instance, was that multiple
conditions on the primeval Earth favoured chemical reactions that synthesized the same set of complex
organic compounds from such simple precursors. Bernal coined the term biopoiesis in 1949 to refer to the
origin of life.[169] In 1967, he suggested that it occurred in three "stages":

1. the origin of biological monomers


2. the origin of biological polymers
3. the evolution from molecules to cells

Bernal suggested that evolution commenced between stages 1 and 2. Bernal regarded the third stage, in
which biological reactions were incorporated behind a cell's boundary, as the most difficult. Modern work
on the way that cell membranes self-assemble, and the work on micropores in various substrates, may be a
key step towards understanding the development of independent free-living cells.[170][171][172]

Miller–Urey experiment

One of the most important pieces of experimental support for the "soup" theory came in 1952. Stanley
Miller and Harold Urey performed an experiment that demonstrated how organic molecules could have
spontaneously formed from inorganic precursors under conditions like those posited by the Oparin-Haldane
hypothesis. The now-famous Miller–Urey experiment used a highly reducing mixture of gases—methane,
ammonia, and hydrogen, as well as water vapor—to form simple organic monomers such as amino
acids.[173] The mixture of gases was cycled through an apparatus that delivered electrical sparks to the
mixture. After one week, it was found that about 10% to 15% of the carbon in the system was then in the
form of a racemic mixture of organic compounds, including amino acids, which are the building blocks of
proteins. This provided direct experimental support for the second point of the "soup" theory, and it is
around the remaining two points of the theory that much of the debate now
centers. A 2011 reanalysis of the saved vials containing the original
extracts that resulted from the Miller and Urey experiments, using current
and more advanced analytical equipment and technology, has uncovered
more biochemicals than originally discovered in the 1950s. One of the
more important findings was 23 amino acids, far more than the five
originally found.[174]

In November 2020, a team of international scientists reported studies


which suggest that the primeval atmosphere of the Earth was much
different than the conditions used in the Miller-Urey studies.[175] Indicated
are results with gas discharge for the origin of life in the primary
atmosphere and hydrosphere.[176]

Stanley Miller
Primordial origin of biological molecules:
Chemistry
The chemical processes on the pre-biotic early Earth are called chemical evolution. The elements, except
for hydrogen and helium, ultimately derive from stellar nucleosynthesis. In 2016, astronomers reported that
the very basic chemical ingredients of life—the carbon-hydrogen molecule (CH, or methylidyne radical),
the carbon-hydrogen positive ion (CH+) and the carbon ion (C+)—are largely the result of ultraviolet light
from stars, rather than other forms of radiation from supernovae and young stars, as thought earlier.[177]
Complex molecules, including organic molecules, form naturally both in space and on planets.[24] There
are two possible sources of organic molecules on the early Earth:

1. Terrestrial origins – organic molecule synthesis driven by impact shocks or by other energy
sources (such as UV light, redox coupling, or electrical discharges; e.g., Miller's
experiments)
2. Extraterrestrial origins – formation of organic molecules in interstellar dust clouds, which rain
down on planets.[178][179] (See pseudo-panspermia)

Observed extraterrestrial organic molecules

An organic compound is any member of a large class of gaseous, liquid, or solid chemicals whose
molecules contain carbon. Carbon is the fourth most abundant element in the Universe by mass after
hydrogen, helium, and oxygen.[180] Carbon is abundant in the Sun, stars, comets, and in the atmospheres
of most planets.[181] Organic compounds are relatively common in space, formed by "factories of complex
molecular synthesis" which occur in molecular clouds and circumstellar envelopes, and chemically evolve
after reactions are initiated mostly by ionizing radiation.[24][182][183][184] Based on computer model
studies, the complex organic molecules necessary for life may have formed on dust grains in the
protoplanetary disk surrounding the Sun before the formation of the Earth.[185] According to the computer
studies, this same process may also occur around other stars that acquire planets.[185]

Amino acids

NASA announced in 2009 that scientists had identified another fundamental chemical building block of life
in a comet for the first time, glycine, an amino acid, which was detected in material ejected from comet
Wild 2 in 2004 and grabbed by NASA's Stardust probe. Glycine has been detected in meteorites before.
Carl Pilcher, who leads the NASA Astrobiology Institute commented that
The discovery of glycine in a comet supports the idea that the fundamental building blocks of
life are prevalent in space, and strengthens the argument that life in the universe may be
common rather than rare.[186]

Comets are encrusted with outer layers of dark material, thought to be a tar-like substance composed of
complex organic material formed from simple carbon compounds after reactions initiated mostly by
ionizing radiation. It is possible that a rain of material from comets could have brought significant quantities
of such complex organic molecules to Earth.[187][188][189] Amino acids which were formed
extraterrestrially may also have arrived on Earth via comets.[104] It is estimated that during the Late Heavy
Bombardment, meteorites may have delivered up to five million tons of organic prebiotic elements to Earth
per year.[104]

PAH world hypothesis

Polycyclic aromatic hydrocarbons (PAH) are the most common and abundant of the known polyatomic
molecules in the observable universe, and are considered a likely constituent of the primordial
sea.[190][191][192] In 2010, PAHs were detected in nebulae.[193]

Polycyclic aromatic hydrocarbons (PAH) are known to


be abundant in the universe,[190][191][192] including in
the interstellar medium, in comets, and in meteorites,
and are some of the most complex molecules so far
found in space.[181]

Other sources of complex molecules have been


The Cat's Paw Nebula lies inside the Milky Way
postulated, including extraterrestrial stellar or interstellar Galaxy and is located in the constellation
origin. For example, from spectral analyses, organic Scorpius.
molecules are known to be present in comets and Green areas show regions where radiation from
meteorites. In 2004, a team detected traces of PAHs in a hot stars collided with large molecules and small
nebula.[194] In 2010, another team also detected PAHs, dust grains called "polycyclic aromatic
along with fullerenes, in nebulae.[193] The use of PAHs hydrocarbons" (PAHs), causing them to
has also been proposed as a precursor to the RNA fluoresce.
world in the PAH world hypothesis. [195] The Spitzer (Spitzer space telescope, 2018)
Space Telescope has detected a star, HH 46-IR,
forming by a process similar to that by which the Sun
formed. In the disk of material surrounding the star, there is a large range of molecules, including cyanide
compounds, hydrocarbons, and carbon monoxide. In 2012, NASA scientists reported that PAHs, subjected
to interstellar medium conditions, are transformed, through hydrogenation, oxygenation and hydroxylation,
to more complex organics—"a step along the path toward amino acids and nucleotides, the raw materials of
proteins and DNA, respectively."[196][197] Further, as a result of these transformations, the PAHs lose their
spectroscopic signature, which could be one of the reasons "for the lack of PAH detection in interstellar ice
grains, particularly the outer regions of cold, dense clouds or the upper molecular layers of protoplanetary
disks."[196][197]

NASA maintains a database for tracking PAHs in the universe.[181][198] More than 20% of the carbon in
the universe may be associated with PAHs,[181] possible starting materials for the formation of life. PAHs
seem to have formed shortly after the Big Bang, are widespread throughout the universe,[190][191][192] and
are associated with new stars and exoplanets.[181]

Nucleobases
Observations suggest that the majority of organic compounds introduced on Earth by interstellar dust
particles are considered principal agents in the formation of complex molecules, thanks to their peculiar
surface-catalytic activities.[199][200] Studies reported in 2008, based on 12 C/13 C isotopic ratios of organic
compounds found in the Murchison meteorite, suggested that the RNA component uracil and related
molecules, including xanthine, were formed extraterrestrially.[201][202] In 2011, a report based on NASA
studies of meteorites found on Earth was published suggesting DNA components (adenine, guanine and
related organic molecules) were made in outer space.[199][203][204] Scientists also found that the cosmic
dust permeating the universe contains complex organics ("amorphous organic solids with a mixed
aromatic–aliphatic structure") that could be created naturally, and rapidly, by stars.[205][206][207] Sun Kwok
of The University of Hong Kong suggested that these compounds may have been related to the
development of life on Earth said that "If this is the case, life on Earth may have had an easier time getting
started as these organics can serve as basic ingredients for life."[205]

The sugar glycolaldehyde

Glycolaldehyde, the first example of an interstellar sugar molecule,


was detected in the star-forming region near the centre of our
galaxy. It was discovered in 2000 by Jes Jørgensen and Jan
Hollis.[208] In 2012, Jørgensen's team reported the detection of
glycolaldehyde in a distant star system. The molecule was found
around the protostellar binary IRAS 16293-2422 400 light years
from Earth.[209][210][211] Glycolaldehyde is needed to form RNA,
which is similar in function to DNA. These findings suggest that
complex organic molecules may form in stellar systems prior to the
formation of planets, eventually arriving on young planets early in
their formation.[212][213] Because sugars are associated with both
metabolism and the genetic code, two of the most basic aspects of
life, it is thought the discovery of extraterrestrial sugar increases the
likelihood that life may exist elsewhere in our galaxy.[208]

Formation of glycolaldehyde in
Polyphosphates stardust

A problem in most scenarios of abiogenesis is that the


thermodynamic equilibrium of amino acid versus peptides is in the direction of separate amino acids. What
has been missing is some force that drives polymerization. The resolution of this problem may well be in
the properties of polyphosphates.[214][215] Polyphosphates are formed by polymerization of ordinary
monophosphate ions PO4 3-. Several mechanisms of organic molecule synthesis have been investigated.
Polyphosphates cause polymerization of amino acids into peptides. They are also logical precursors in the
synthesis of such key biochemical compounds as adenosine triphosphate (ATP). A key issue seems to be
that calcium reacts with soluble phosphate to form insoluble calcium phosphate (apatite), so some plausible
mechanism must be found to keep calcium ions from causing precipitation of phosphate. There has been
much work on this topic over the years, but an interesting new idea is that meteorites may have introduced
reactive phosphorus species on the early Earth.[216] Based on recent computer model studies, the complex
organic molecules necessary for life may have formed in the protoplanetary disk of dust grains surrounding
the Sun before the formation of the Earth.[185][217] According to the computer studies, this same process
may also occur around other stars that acquire planets. (Also see Extraterrestrial organic molecules).

The accumulation and concentration of organic molecules on a planetary surface is also considered an
essential early step for the origin of life.[44] Identifying and understanding the mechanisms that led to the
production of prebiotic molecules in various environments is critical for establishing the inventory of
ingredients from which life originated on Earth, assuming that the abiotic production of molecules
ultimately influenced the selection of molecules from which life emerged.[44]

In 2019, scientists reported detecting, for the first time, sugar molecules, including ribose, in meteorites,
suggesting that chemical processes on asteroids can produce some fundamentally essential bio-ingredients
important to life, and supporting the notion of an RNA world prior to a DNA-based origin of life on Earth,
and possibly, as well, the notion of panspermia.[218][213]

Chemical synthesis in the laboratory

As early as the 1860s, experiments have demonstrated that biologically relevant molecules can be produced
from interaction of simple carbon sources with abundant inorganic catalysts.

Fox proteinoids

In trying to uncover the intermediate stages of abiogenesis mentioned by Bernal, Sidney Fox in the 1950s
and 1960s studied the spontaneous formation of peptide structures (small chains of amino acids) under
conditions that might plausibly have existed early in Earth's history. In one of his experiments, he allowed
amino acids to dry out as if puddled in a warm, dry spot in prebiotic conditions: In an experiment to set
suitable conditions for life to form, Fox collected volcanic material from a cinder cone in Hawaii. He
discovered that the temperature was over 100 C just 4 inches (100 mm) beneath the surface of the cinder
cone, and suggested that this might have been the environment in which life was created—molecules could
have formed and then been washed through the loose volcanic ash into the sea. He placed lumps of lava
over amino acids derived from methane, ammonia and water, sterilized all materials, and baked the lava
over the amino acids for a few hours in a glass oven. A brown, sticky substance formed over the surface,
and when the lava was drenched in sterilized water, a thick, brown liquid leached out. He found that, as
they dried, the amino acids formed long, often cross-linked, thread-like, submicroscopic polypeptide
molecules.[219]

Sugars

In particular, experiments by Butlerov (the formose reaction) showed that tetroses, pentoses, and hexoses
are produced when formaldehyde is heated under basic conditions with divalent metal ions like calcium.
The reaction was scrutinized and subsequently proposed to be autocatalytic by Breslow in 1959.

Nucleobases

Similar experiments (see below) demonstrate that nucleobases like guanine and adenine could be
synthesized from simple carbon and nitrogen sources like hydrogen cyanide and ammonia.

Formamide produces all four ribonucleotides and other biological molecules when warmed in the presence
of various terrestrial minerals. Formamide is ubiquitous in the Universe, produced by the reaction of water
and hydrogen cyanide (HCN). It has several advantages as a biotic precursor, including the ability to easily
become concentrated through the evaporation of water.[220][221] Although HCN is poisonous, it only
affects aerobic organisms (eukaryotes and aerobic bacteria), which did not yet exist. It can play roles in
other chemical processes as well, such as the synthesis of the amino acid glycine.[104]

In March 2015, NASA scientists reported that, for the first time, complex DNA and RNA organic
compounds of life, including uracil, cytosine and thymine, have been formed in the laboratory under outer
space conditions, using starting chemicals, such as pyrimidine, found in meteorites. Pyrimidine, like PAHs,
the most carbon-rich chemical found in the Universe, may have been formed in red giant stars or in
interstellar dust and gas clouds.[222] A group of Czech scientists reported that all four RNA-bases may be
synthesized from formamide in the course of high-energy density events like extraterrestrial impacts.[223]

Use of reactions for ammonium cyanide

In 1961, it was shown that the nucleic acid purine base adenine can be formed by heating aqueous
ammonium cyanide solutions.[224] The boiling point of ammonium cyanide is 36°С.

Effects with temperatures around the freezing point of water

Other pathways for synthesizing bases from inorganic materials were also reported.[225] Orgel and
colleagues have shown that freezing temperatures are advantageous for the synthesis of purines, due to the
concentrating effect for key precursors such as hydrogen cyanide.[226] Research by Miller and colleagues
suggested that while adenine and guanine require freezing conditions for synthesis, cytosine and uracil may
require boiling temperatures.[227] Research by the Miller group notes the formation of seven different
amino acids and 11 types of nucleobases in ice when ammonia and cyanide were left in a freezer from 1972
to 1997.[228][229] Other work demonstrated the formation of s-triazines (alternative nucleobases),
pyrimidines (including cytosine and uracil), and adenine from urea solutions subjected to freeze-thaw
cycles under a reductive atmosphere (with spark discharges as an energy source).[230] The explanation
given for the unusual speed of these reactions at such a low temperature is eutectic freezing. As an ice
crystal forms, it stays pure: only molecules of water join the growing crystal, while impurities like salt or
cyanide are excluded. These impurities become crowded in microscopic pockets of liquid within the ice,
and this crowding causes the molecules to collide more often. Mechanistic exploration using quantum
chemical methods provide a more detailed understanding of some of the chemical processes involved in
chemical evolution, and a partial answer to the fundamental question of molecular biogenesis.[231]

Use of less-reducing gas in Miller–Urey experiment

At the time of the Miller–Urey experiment, scientific consensus was that the early Earth had a reducing
atmosphere with compounds relatively rich in hydrogen and poor in oxygen (e.g., CH4 and NH3 as
opposed to CO2 and nitrogen dioxide (NO2 )). However, current scientific consensus describes the
primitive atmosphere as either weakly reducing or neutral[232][233] (see also Oxygen Catastrophe). Such an
atmosphere would diminish both the amount and variety of amino acids that could be produced, although
studies that include iron and carbonate minerals (thought present in early oceans) in the experimental
conditions have again produced a diverse array of amino acids.[232] Other scientific research has focused
on two other potential reducing environments: outer space and deep-sea thermal vents.[234][235][236]

Synthesis based on hydrogen cyanide

A research project completed in 2015 by John Sutherland and others found that a network of reactions
beginning with hydrogen cyanide and hydrogen sulfide, in streams of water irradiated by UV light, could
produce the chemical components of proteins and lipids, as well as those of RNA,[237][238] while not
producing a wide range of other compounds.[239] The researchers used the term "cyanosulfidic" to describe
this network of reactions.[238]

Issues during laboratory synthesis


The spontaneous formation of complex polymers from abiotically generated monomers under the
conditions posited by the "soup" theory is not at all a straightforward process. Besides the necessary basic
organic monomers, compounds that would have prohibited the formation of polymers were also formed in
high concentration during the Miller–Urey and Joan Oró experiments.[240] The Miller–Urey experiment,
for example, produces many substances that would react with the amino acids or terminate their coupling
into peptide chains.[241] Biology uses essentially 20 amino acids for its coded protein enzymes,
representing a very small subset of the structurally possible set. Most models of the origin of life suggest
organisms developed from environmentally available organic compounds.[242] The fundamental roles that
peptides and proteins play in today's biology makes it almost indisputable that peptides were key players in
the origin of life. Insofar as it is appropriate to extrapolate back from extant biology to the prebiotic world,
one must acknowledge the critical importance that interconnected molecular networks, likely with peptides
as key components, would have played in life's origin.[243]

Autocatalysis

Autocatalysts are substances that catalyze the production of themselves and therefore are "molecular
replicators." The simplest self-replicating chemical systems are autocatalytic, and typically contain three
components: a product molecule and two precursor molecules. The product molecule joins the precursor
molecules, which in turn produce more product molecules from more precursor molecules. The product
molecule catalyzes the reaction by providing a complementary template that binds to the precursors, thus
bringing them together. Such systems have been demonstrated both in biological macromolecules and in
small organic molecules.[244][245] Systems that do not proceed by template mechanisms, such as the self-
reproduction of micelles and vesicles, have also been observed.[245]

It has been proposed that life initially arose as autocatalytic chemical networks.[246] British ethologist
Richard Dawkins wrote about autocatalysis as a potential explanation for the origin of life in his 2004 book
The Ancestor's Tale.[247] In his book, Dawkins cites experiments performed by Julius Rebek and his
colleagues in which they combined amino adenosine and pentafluorophenyl esters with the autocatalyst
amino adenosine triacid ester (AATE). One product was a variant of AATE, which catalyzed the synthesis
of themselves. This experiment demonstrated the possibility that autocatalysts could exhibit competition
within a population of entities with heredity, which could be interpreted as a rudimentary form of natural
selection.[248][249]

Pertinent geological environments

Darwin's little pond

An early concept, that life originated from non-living matter in slow stages, appeared in Herbert Spencer's
1864–1867 book Principles of Biology. In 1879 William Turner Thiselton-Dyer referred to this in a paper
"On spontaneous generation and evolution". On 1 February 1871 Charles Darwin wrote about these
publications to Joseph Hooker, and set out his own speculation,[250][251][252] suggesting that the original
spark of life may have begun in a

warm little pond, with all sorts of ammonia and phosphoric salts, light, heat, electricity, &c.,
present, that a proteine compound was chemically formed ready to undergo still more complex
changes.

He went on to explain that


at the present day such matter would be instantly devoured or absorbed, which would not have
been the case before living creatures were formed.

Darwin 1887, p. 18 (http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F1452.3&pages


eq=30):

More recent studies, in 2017, support the notion that life may have begun right after the Earth was formed
as RNA molecules emerging from "warm little ponds".[253]

Volcanic hot springs and hydrothermal vents, shallow or deep

Martin Brazier has shown that early micro-fossils came from a hot world of gases such as methane,
ammonia, carbon dioxide and hydrogen sulphide, which are toxic to much current life.[254] Another
analysis of the conventional threefold tree of life shows thermophilic and hyperthermophilic bacteria and
archaea are closest to the root, suggesting that life may have evolved in a hot environment.[255]

Deep sea hydrothermal vents

The deep sea vent, or alkaline hydrothermal vent, theory posits that life
may have begun at submarine hydrothermal vents,[256][257] Martin and
Russell have suggested

that life evolved in structured iron monosulphide precipitates


in a seepage site hydrothermal mound at a redox, pH, and
temperature gradient between sulphide-rich hydrothermal
fluid and iron(II)-containing waters of the Hadean ocean floor.
The naturally arising, three-dimensional compartmentation
observed within fossilized seepage-site metal sulphide
precipitates indicates that these inorganic compartments were
the precursors of cell walls and membranes found in free-
living prokaryotes. The known capability of FeS and NiS to Deep-sea hydrothermal vent
catalyze the synthesis of the acetyl-methylsulphide from or black smoker
carbon monoxide and methylsulphide, constituents of
hydrothermal fluid, indicates that pre-biotic syntheses
occurred at the inner surfaces of these metal-sulphide-walled
compartments,...[258]

These form where hydrogen-rich fluids emerge from below the sea floor, as a result of serpentinization of
ultra-mafic olivine with seawater and a pH interface with carbon dioxide-rich ocean water. The vents form
a sustained chemical energy source derived from redox reactions, in which electron donors (molecular
hydrogen) react with electron acceptors (carbon dioxide); see Iron–sulfur world theory. These are
exothermic reactions.[256][b]

Russell demonstrated that alkaline vents created an abiogenic proton motive force (PMF) chemiosmotic
gradient,[258] in which conditions are ideal for an abiogenic hatchery for life. Their microscopic
compartments "provide a natural means of concentrating organic molecules," composed of iron-sulfur
minerals such as mackinawite, endowed these mineral cells with the catalytic properties envisaged by
Günter Wächtershäuser.[259] This movement of ions across the membrane depends on a combination of
two factors:
1. Diffusion force caused by concentration gradient—all particles including ions tend to diffuse
from higher concentration to lower.
2. Electrostatic force caused by electrical potential gradient—cations like protons H+ tend to
diffuse down the electrical potential, anions in the opposite direction.

These two gradients taken together can be expressed as an electrochemical gradient, providing energy for
abiogenic synthesis. The proton motive force can be described as the measure of the potential energy stored
as a combination of proton and voltage gradients across a membrane (differences in proton concentration
and electrical potential).

Szostak suggested that geothermal activity provides greater opportunities for the origination of life in open
lakes where there is a buildup of minerals. In 2010, based on spectral analysis of sea and hot mineral water,
Ignat Ignatov and Oleg Mosin demonstrated that life may have predominantly originated in hot mineral
water. The hot mineral water that contains hydrocarbonate and calcium ions has the most optimal
range.[260][261] This case is similar to the origin of life in hydrothermal vents, but with hydrocarbonate and
calcium ions in hot water. The research with spectral analyses was performed in Rupite, Bulgaria, with hot
mineral water with calcium and hydrocarbonate ions, Anoxybacillus rupiences sp., bacteria, archaea and
cyanobacteria [262][263] Mineral water with pH of 9–11 is possible to have the reactions in seawater.
According to Melvin Calvin, certain reactions of condensation-dehydration of amino acids and nucleotides
in individual blocks of peptides and nucleic acids can take place in the primary hydrosphere with pH 9–11
at a later evolutionary stage.[264] Some of these compounds like hydrocyanic acid (HCN) have been
proven in the experiments of Miller. This is the environment in which the stromatolites have been created.
David Ward of Montana State University described the formation of stromatolites in hot mineral water at
the Yellowstone National Park. Stromatolites survive in hot mineral water and in proximity to areas with
volcanic activity.[265] Processes have evolved in the sea near geysers of hot mineral water. In 2011, Tadashi
Sugawara from the University of Tokyo created a protocell in hot water.[266]

Experimental research and computer modeling suggest that the surfaces of mineral particles inside
hydrothermal vents have catalytic properties similar to those of enzymes and are able to create simple
organic molecules, such as methanol (CH3 OH) and formic, acetic and pyruvic acid out of the dissolved
CO2 in the water if driven by an applied voltage or by reaction with H2 or H2 S.[267][268]

The research reported above by Martin in 2016 supports the thesis that life arose at hydrothermal
vents,[269][270] that spontaneous chemistry in the Earth's crust driven by rock–water interactions at
disequilibrium thermodynamically underpinned life's origin[271][272] and that the founding lineages of the
archaea and bacteria were H2 -dependent autotrophs that used CO2 as their terminal acceptor in energy
metabolism.[273] Martin suggests, based upon this evidence that LUCA "may have depended heavily on
the geothermal energy of the vent to survive".[274]

Fluctuating hydrothermal pools on volcanic islands or proto-continents

Mulkidjanian and co-authors think that the marine environments did not provide the ionic balance and
composition universally found in cells, as well as of ions required by essential proteins and ribozymes
found in virtually all living organisms, especially with respect to K+/Na+ ratio, Mn2+, Zn2+ and phosphate
concentrations. The only known environments that mimic the needed conditions on Earth are found in
terrestrial hydrothermal pools fed by steam vents.[256] Additionally, mineral deposits in these environments
under an anoxic atmosphere would have suitable pH (as opposed to current pools in an oxygenated
atmosphere), contain precipitates of sulfide minerals that block harmful UV radiation, have wetting/drying
cycles that concentrate substrate solutions to concentrations amenable to spontaneous formation of
polymers of nucleic acids, polyesters[275] and depsipeptides,[276] both by chemical reactions in the
hydrothermal environment, as well as by exposure to UV light during transport from vents to adjacent
pools. Their hypothesized pre-biotic environments are similar to the deep-oceanic vent environments most
commonly hypothesized, but add additional components that help explain peculiarities found in
reconstructions of the Last Universal Common Ancestor (LUCA) of all living organisms.[277]

Colín-García et al. (2016) discuss the advantages and disadvantages of hydrothermal vents as primitive
environments.[256] They mention the exergonic reactions in such systems could have been a source of free
energy that promoted chemical reactions, additional to their high mineralogical diversity which implies the
induction of important chemical gradients, thus favoring the interaction between electron donors and
acceptors. Colín-García et al. (2016) also summarize a set of experiments proposed to test the role of
hydrothermal vents in prebiotic synthesis.[256]

Volcanic ash in the ocean

Geoffrey W. Hoffmann has argued that a complex nucleation event as the origin of life involving both
polypeptides and nucleic acid is compatible with the time and space available in the primary oceans of
Earth[278] Hoffmann suggests that volcanic ash may provide the many random shapes needed in the
postulated complex nucleation event. This aspect of the theory can be tested experimentally.

Gold's deep-hot biosphere

In the 1970s, Thomas Gold proposed the theory that life first developed not on the surface of the Earth, but
several kilometers below the surface. It is claimed that the discovery of microbial life below the surface of
another body in our Solar System would lend significant credence to this theory. Gold also asserted that a
trickle of food from a deep, unreachable, source is needed for survival because life arising in a puddle of
organic material is likely to consume all of its food and become extinct. Gold's theory is that the flow of
such food is due to out-gassing of primordial methane from the Earth's mantle; more conventional
explanations of the food supply of deep microbes (away from sedimentary carbon compounds) is that the
organisms subsist on hydrogen released by an interaction between water and (reduced) iron compounds in
rocks.

Radioactive beach hypothesis

Zachary Adam claims that tidal processes that occurred during a time when the Moon was much closer
may have concentrated grains of uranium and other radioactive elements at the high-water mark on
primordial beaches, where they may have been responsible for generating life's building blocks.[279]
According to computer models,[280] a deposit of such radioactive materials could show the same self-
sustaining nuclear reaction as that found in the Oklo uranium ore seam in Gabon. Such radioactive beach
sand might have provided sufficient energy to generate organic molecules, such as amino acids and sugars
from acetonitrile in water. Radioactive monazite material also has released soluble phosphate into the
regions between sand-grains, making it biologically "accessible." Thus amino acids, sugars, and soluble
phosphates might have been produced simultaneously, according to Adam. Radioactive actinides, left
behind in some concentration by the reaction, might have formed part of organometallic complexes. These
complexes could have been important early catalysts to living processes.

John Parnell has suggested that such a process could provide part of the "crucible of life" in the early stages
of any early wet rocky planet, so long as the planet is large enough to have generated a system of plate
tectonics which brings radioactive minerals to the surface. As the early Earth is thought to have had many
smaller plates, it might have provided a suitable environment for such processes.[281]
Thermodynamics, self-organization, and information: Physics

Thermodynamics principles: Energy and entropy

In antiquity it was commonly thought, for instance by Empedocles and Aristotle, that the life of the
individuals of some species, and more generally, life itself, could start with high temperature, i.e. implicitly
by thermal cycling.[282]

Similarly, it was realized early on that life requires a loss of entropy, or disorder, when molecules organize
themselves into living matter. This Second Law of thermodynamics needs to be considered when self-
organization of matter to higher complexity happens. Because living organisms are machines,[283] the
Second Law applies to life as well.

Thus, contrary to a naive view of the Second Law, the emergence of life and increased complexity does not
contradict this law: First, a living organism creates order in some places (e.g. its living body or dwelling) at
the expense of an increase of entropy elsewhere (e.g. heat and waste production). Second, the Second Law
of thermodynamics actually predicts an increase in complexity[284] and in correlations between a system
and its surrounding, when undergoing interaction[285] - with memory and genetic adaptation being
examples of such correlations between a living organism and its environment.

Obtaining free energy

Bernal said on the Miller–Urey experiment that

it is not enough to explain the formation of such molecules, what is necessary, is a physical-
chemical explanation of the origins of these molecules that suggests the presence of suitable
sources and sinks for free energy.[286]

Multiple sources of energy were available for chemical reactions on the early Earth. For example, heat
(such as from geothermal processes) is a standard energy source for chemistry. Other examples include
sunlight and electrical discharges (lightning), among others.[104] In fact, lightning is a plausible energy
source for the origin of life, given that just in the tropics lightning strikes about 100 million times a
year.[287]

Computer simulations also suggest that cavitation in primordial water reservoirs such as breaking sea
waves, streams and oceans can potentially lead to the synthesis of biogenic compounds.[288]

Unfavorable reactions can also be driven by highly favorable ones, as in the case of iron-sulfur chemistry.
For example, this was probably important for carbon fixation (the conversion of carbon from its inorganic
form to an organic one).[c] Carbon fixation by reaction of CO2 with H2 S via iron-sulfur chemistry is
favorable, and occurs at neutral pH and 100 °C. Iron-sulfur surfaces, which are abundant near
hydrothermal vents, are also capable of producing small amounts of amino acids and other biological
metabolites.[104]

Self-organization
The discipline of synergetics studies self-organization in physical systems.
In his book Synergetics[289] Hermann Haken has pointed out that different
physical systems can be treated in a similar way. He gives as examples of
self-organization several types of lasers, instabilities in fluid dynamics,
including convection, and chemical and biochemical oscillations. In his
preface he mentions the origin of life, but only in general terms:

The spontaneous formation of well organized structures out of


germs or even out of chaos is one of the most fascinating
phenomena and most challenging problems scientists are
confronted with. Such phenomena are an experience of our Hermann Haken
daily life when we observe the growth of plants and animals.
Thinking of much larger time scales, scientists are led into the
problems of evolution, and, ultimately, of the origin of living
matter. When we try to explain or understand in some sense
these extremely complex biological phenomena it is a natural
question, whether processes of self-organization may be
found in much simpler systems of the unanimated world.

In recent years it has become more and more evident that


there exists numerous examples in physical and chemical
systems where well organized spatial, temporal, or spatio-
temporal structures arise out of chaotic states. Furthermore, as
in living organisms, the functioning of these systems can be
maintained only by a flux of energy (and matter) through
them. In contrast to man-made machines, which are devised to
exhibit special structures and functionings, these structures
develop spontaneously—they are selforganizing. ...[290]

Dissipative structuring

This theory postulates that the hallmark of the origin and evolution of life is the microscopic dissipative
structuring under UVC light of organic pigments and their proliferation over the entire Earth
surface.[291][292][293] Present day life augments the entropy production of Earth in its solar environment by
dissipating ultraviolet and visible photons into heat through organic pigments in water. This heat then
catalyzes a host of secondary dissipative processes such as the water cycle, ocean and wind currents,
hurricanes, etc.[294]

Self-organization by dissipative structures

The 19th-century physicist Ludwig Boltzmann first recognized that the struggle for existence of living
organisms was neither over raw material nor energy, but instead had to do with entropy production derived
from the conversion of the solar spectrum into heat by these systems.[295] Boltzmann thus realized that
living systems, like all irreversible processes, were dependent on the dissipation of a generalized chemical
potential for their existence. In his book "What is Life", the 20th-century physicist Erwin Schrödinger[296]
emphasized the importance of Boltzmann's deep insight into the irreversible thermodynamic nature of living
systems, suggesting that this was the physics and chemistry behind the origin and evolution of life.
However, irreversible processes, and much less living systems, could not
be conveniently analyzed under this perspective until Lars Onsager,[297]
and later Ilya Prigogine,[298] developed an elegant mathematical formalism
for treating the "self-organization" of material under a generalized
chemical potential. This formalism became known as Classical Irreversible
Thermodynamics and Prigogine was awarded the Nobel Prize in
Chemistry in 1977 "for his contributions to non-equilibrium
thermodynamics, particularly the theory of dissipative structures". The
analysis by Prigogine showed that if a system were left to evolve under an
imposed external potential, material could spontaneously organize (lower
its entropy) forming what he called "dissipative structures" which would
increase the dissipation of the externally imposed potential (augment the
global entropy production). Non-equilibrium thermodynamics has since
Ilya Prigogine 1977c
been successfully applied to the analysis of living systems, from the
biochemical production of ATP[299] to optimizing bacterial metabolic
pathways[300] to complete ecosystems.[301][302][303]

Encapsulation: morphology

Encapsulation without a membrane

Oparin's coacervate

Membraneless polyester droplets

Researchers Tony Jia and Kuhan Chandru[304] have proposed that membraneless polyesters droplets could
have been significant in the Origins of Life.[305] Given the "messy" nature of prebiotic chemistry,[306][307]
the spontaneous generation of these combinatorial droplets may have played a role in early cellularization
before the innovation of lipid vesicles. Protein function within and RNA function in the presence of certain
polyester droplets was shown to be preserved within the droplets. Additionally, the droplets have
scaffolding ability, by allowing lipids to assemble around them that may have prevented leakage of genetic
materials.

Proteinoid microspheres

Fox observed in the 1960s that the proteinoids that he had synthesized could form cell-like structures that
have been named "proteinoid microspheres".[219]

The amino acids had combined to form proteinoids, and the proteinoids had combined to form small
globules that Fox called "microspheres". His proteinoids were not cells, although they formed clumps and
chains reminiscent of cyanobacteria, but they contained no functional nucleic acids or any encoded
information. Based upon such experiments, Colin Pittendrigh stated in 1967 that "laboratories will be
creating a living cell within ten years," a remark that reflected the typical contemporary naivety about the
complexity of cell structures.[308]

Lipid world
The lipid world theory postulates that the first self-replicating object was lipid-like.[309][310] It is known
that phospholipids form lipid bilayers in water while under agitation—the same structure as in cell
membranes. These molecules were not present on early Earth, but other amphiphilic long-chain molecules
also form membranes. Furthermore, these bodies may expand (by insertion of additional lipids), and under
excessive expansion may undergo spontaneous splitting which preserves the same size and composition of
lipids in the two progenies. The main idea in this theory is that the molecular composition of the lipid
bodies is the preliminary way for information storage, and evolution led to the appearance of polymer
entities such as RNA or DNA that may store information favourably. Studies on vesicles from potentially
prebiotic amphiphiles have so far been limited to systems containing one or two types of amphiphiles. This
in contrast to the output of simulated prebiotic chemical reactions, which typically produce very
heterogeneous mixtures of compounds.[311] Within the hypothesis of a lipid bilayer membrane composed
of a mixture of various distinct amphiphilic compounds there is the opportunity of a huge number of
theoretically possible combinations in the arrangements of these amphiphiles in the membrane. Among all
these potential combinations, a specific local arrangement of the membrane would have favoured the
constitution of a hypercycle,[312][313] actually a positive feedback composed of two mutual catalysts
represented by a membrane site and a specific compound trapped in the vesicle. Such site/compound pairs
are transmissible to the daughter vesicles leading to the emergence of distinct lineages of vesicles which
would have allowed Darwinian natural selection.[314]

Protocells

A protocell is a self-organized, self-ordered, spherical collection of lipids


proposed as a stepping-stone to the origin of life.[311] A central question in
evolution is how simple protocells first arose and differed in reproductive
contribution to the following generation driving the evolution of life.
Although a functional protocell has not yet been achieved in a laboratory
setting, there are scientists who think the goal is well within
reach.[315][316][317]

Self-assembled vesicles are essential components of primitive cells.[311]


The second law of thermodynamics requires that the universe move in a
direction in which entropy increases, yet life is distinguished by its great
degree of organization. Therefore, a boundary is needed to separate life The three main structures
processes from non-living matter. [318] Researchers Irene Chen and Szostak phospholipids form
amongst others, suggest that simple physicochemical properties of spontaneously in solution:
elementary protocells can give rise to essential cellular behaviours, the liposome (a closed
bilayer), the micelle and the
including primitive forms of differential reproduction competition and
bilayer.
energy storage. Such cooperative interactions between the membrane and
its encapsulated contents could greatly simplify the transition from simple
replicating molecules to true cells.[316] Furthermore, competition for
membrane molecules would favour stabilized membranes, suggesting a selective advantage for the
evolution of cross-linked fatty acids and even the phospholipids of today.[316] Such micro-encapsulation
would allow for metabolism within the membrane, the exchange of small molecules but the prevention of
passage of large substances across it.[319] The main advantages of encapsulation include the increased
solubility of the contained cargo within the capsule and the storage of energy in the form of an
electrochemical gradient.

Another approach to the notion of a protocell concerns the term "chemoton" (short for 'chemical
automaton') which refers to an abstract model for the fundamental unit of life introduced by Hungarian
theoretical biologist Tibor Gánti.[51] It is the oldest known computational abstract of a protocell. Gánti
conceived the basic idea in 1952 and formulated the concept in 1971 in his book The Principles of Life
(originally written in Hungarian, and translated to English only in 2003). He surmised the chemoton as the
original ancestor of all organisms, or the last universal common ancestor.[320]

The basic assumption of the chemoton model is that life should fundamentally and essentially have three
properties: metabolism, self-replication, and a bilipid membrane.[321] The metabolic and replication
functions together form an autocatalytic subsystem necessary for the basic functions of life, and a
membrane encloses this subsystem to separate it from the surrounding environment. Therefore, any system
having such properties may be regarded as alive, and it will be subjected to natural selection and contain a
self-sustaining cellular information. Some consider this model a significant contribution to origin of life as it
provides a philosophy of evolutionary units.[322]

Nonetheless, a 2012 study led by Mulkidjanian of the University of Osnabrück, suggests that inland pools
of condensed and cooled geothermal vapor have the ideal characteristics for the origin of life.[323] Scientists
confirmed in 2002 that by adding a montmorillonite clay to a solution of fatty acid micelles (lipid spheres),
the clay sped up the rate of vesicles formation 100-fold.[317] Furthermore, recent studies have found that
the repeated actions of dehydration and rehydration trapped biomolecules like RNA inside the lipid
protocells found within hot springs and providing the necessary preconditions for evolution by natural
selection.[324]

Lipid vesicles formation in fresh water

Bruce Damer and David Deamer have come to the conclusion that cell membranes cannot be formed in
salty seawater, and must therefore have originated in freshwater. Before the continents formed, the only dry
land on Earth would be volcanic islands, where rainwater would form ponds where lipids could form the
first stages towards cell membranes. These predecessors of true cells are assumed to have behaved more
like a superorganism rather than individual structures, where the porous membranes would house molecules
which would leak out and enter other protocells. Only when true cells had evolved would they gradually
adapt to saltier environments and enter the ocean.[325]

Vesicles consisting of mixtures of RNA-like biochemicals

Another protocell model is the Jeewanu. First synthesized in 1963 from simple minerals and basic organics
while exposed to sunlight, it is still reported to have some metabolic capabilities, the presence of
semipermeable membrane, amino acids, phospholipids, carbohydrates and RNA-like molecules.[326][327]
However, the nature and properties of the Jeewanu remains to be clarified.

Electrostatic interactions induced by short, positively charged, hydrophobic peptides containing 7 amino
acids in length or fewer, can attach RNA to a vesicle membrane, the basic cell membrane.[328][329]

Metal-sulfide precipitates

William Martin and Michael Russell have suggested

. . . . that life evolved in structured iron monosulphide precipitates in a seepage site


hydrothermal mound at a redox, pH, and temperature gradient between sulphide-rich
hydrothermal fluid and iron(II)-containing waters of the Hadean ocean floor. The naturally
arising, three-dimensional compartmentation observed within fossilized seepage-site metal
sulphide precipitates indicates that these inorganic compartments were the precursors of cell
walls and membranes found in free-living prokaryotes. The known capability of FeS and NiS
to catalyze the synthesis of the acetyl-methylsulphide from carbon monoxide and
methylsulphide, constituents of hydrothermal fluid, indicates that pre-biotic syntheses occurred
at the inner surfaces of these metal-sulphide-walled compartments,..."[258]

Origin of metabolism: physiology


Different forms of life with variable origin processes may have appeared quasi-simultaneously in the early
history of Earth.[330] The other forms may be extinct (having left distinctive fossils through their different
biochemistry—e.g., hypothetical types of biochemistry). It has been proposed that:

The first organisms were self-replicating iron-rich clays which fixed carbon dioxide into oxalic
and other dicarboxylic acids. This system of replicating clays and their metabolic phenotype
then evolved into the sulfide rich region of the hotspring acquiring the ability to fix nitrogen.
Finally phosphate was incorporated into the evolving system which allowed the synthesis of
nucleotides and phospholipids. If biosynthesis recapitulates biopoiesis, then the synthesis of
amino acids preceded the synthesis of the purine and pyrimidine bases. Furthermore, the
polymerization of the amino acid thioesters into polypeptides preceded the directed
polymerization of amino acid esters by polynucleotides.[331]

Metabolism-like reactions could have occurred naturally in early oceans, before the first organisms
evolved.[20][332] Metabolism may predate the origin of life, which may have evolved from the chemical
conditions in the earliest oceans. Reconstructions in laboratories show that some of these reactions can
produce RNA, and some others resemble two essential reaction cascades of metabolism: glycolysis and the
pentose phosphate pathway, that provide essential precursors for nucleic acids, amino acids and lipids.[332]

Clay hypothesis

Montmorillonite, an abundant clay, is a catalyst for the polymerization of RNA and for the formation of
membranes from lipids.[333] A model for the origin of life using clay was forwarded by Alexander Cairns-
Smith in 1985 and explored as a plausible mechanism by several scientists.[334] The clay hypothesis
postulates that complex organic molecules arose gradually on pre-existing, non-organic replication surfaces
of silicate crystals in solution.

At the Rensselaer Polytechnic Institute, James Ferris' studies have also confirmed that montmorillonite clay
minerals catalyze the formation of RNA in aqueous solution, by joining nucleotides to form longer
chains.[335]

In 2007, Bart Kahr from the University of Washington and colleagues reported their experiments that tested
the idea that crystals can act as a source of transferable information, using crystals of potassium hydrogen
phthalate. "Mother" crystals with imperfections were cleaved and used as seeds to grow "daughter" crystals
from solution. They then examined the distribution of imperfections in the new crystals and found that the
imperfections in the mother crystals were reproduced in the daughters, but the daughter crystals also had
many additional imperfections. For gene-like behavior to be observed, the quantity of inheritance of these
imperfections should have exceeded that of the mutations in the successive generations, but it did not. Thus
Kahr concluded that the crystals "were not faithful enough to store and transfer information from one
generation to the next."[336]
Iron–sulfur world

In the 1980s, Günter Wächtershäuser, encouraged and supported by Karl Popper,[337][338][339] postulated
his iron–sulfur world, a theory of the evolution of pre-biotic chemical pathways as the starting point in the
evolution of life. It systematically traces today's biochemistry to primordial reactions which provide
alternative pathways to the synthesis of organic building blocks from simple gaseous compounds.

In contrast to the classical Miller experiments, which depend on external sources of energy (simulated
lightning, ultraviolet irradiation), "Wächtershäuser systems" come with a built-in source of energy: sulfides
of iron (iron pyrite) and other minerals. The energy released from redox reactions of these metal sulfides is
available for the synthesis of organic molecules, and such systems may have evolved into autocatalytic sets
constituting self-replicating, metabolically active entities predating the life forms known today.[20][332]
Experiments with such sulfides in an aqueous environment at 100 °C produced a relatively small yield of
dipeptides (0.4% to 12.4%) and a smaller yield of tripeptides (0.10%) although under the same conditions,
dipeptides were quickly broken down.[340]

Several models reject the self-replication of a "naked-gene", postulating instead the emergence of a
primitive metabolism providing a safe environment for the later emergence of RNA replication. The
centrality of the Krebs cycle (citric acid cycle) to energy production in aerobic organisms, and in drawing in
carbon dioxide and hydrogen ions in biosynthesis of complex organic chemicals, suggests that it was one of
the first parts of the metabolism to evolve.[259] Concordantly, geochemists Szostak and Adamala
demonstrated non enzymatic RNA replication in primitive protocells is only possibly in presence of weak
cation chelator like citric acid, providing further evidence for central role of citric acid in primordial
metabolism. doi:10.1126/science.1241888 (https://doi.org/10.1126%2Fscience.1241888)

Russell has proposed that "the purpose of life is to hydrogenate carbon dioxide" (as part of a "metabolism-
first," rather than a "genetics-first," scenario).[341][342] Physicist Jeremy England has proposed that life was
inevitable from general thermodynamic considerations:

... when a group of atoms is driven by an external source of energy (like the sun or chemical
fuel) and surrounded by a heat bath (like the ocean or atmosphere), it will often gradually
restructure itself in order to dissipate increasingly more energy. This could mean that under
certain conditions, matter inexorably acquires the key physical attribute associated with
life.[343][344]

One of the earliest incarnations of this idea was put forward in 1924 with Oparin's notion of primitive self-
replicating vesicles which predated the discovery of the structure of DNA. Variants in the 1980s and 1990s
include Wächtershäuser's iron–sulfur world theory and models introduced by Christian de Duve based on
the chemistry of thioesters. More abstract and theoretical arguments for the plausibility of the emergence of
metabolism without the presence of genes include a mathematical model introduced by Freeman Dyson in
the early 1980s and Stuart Kauffman's notion of collectively autocatalytic sets, discussed later that decade.

Orgel summarized his analysis by stating,

There is at present no reason to expect that multistep cycles such as the reductive citric acid
cycle will self-organize on the surface of FeS/FeS2 or some other mineral."[345]
It is possible that another type of metabolic pathway was used at the beginning of life. For example, instead
of the reductive citric acid cycle, the "open" acetyl-CoA pathway (another one of the five recognized ways
of carbon dioxide fixation in nature today) would be compatible with the idea of self-organization on a
metal sulfide surface. The key enzyme of this pathway, carbon monoxide dehydrogenase/acetyl-CoA
synthase, harbors mixed nickel-iron-sulfur clusters in its reaction centers and catalyzes the formation of
acetyl-CoA (similar to acetyl-thiol) in a single step. There are increasing concerns, however, that prebiotic
thiolated and thioester compounds are thermodynamically and kinetically unfavorable to accumulate in
presumed prebiotic conditions (i.e. hydrothermal vents).[346] It has also been proposed that cysteine and
homocysteine may have reacted with nitriles resulting from the Stecker reaction, readily forming catalytic
thiol-rich poplypeptides.[347]

Zinc-world hypothesis

The zinc world (Zn-world) hypothesis of Mulkidjanian [348] is an extension of Wächtershäuser's pyrite
hypothesis. Wächtershäuser based his theory of the initial chemical processes leading to informational
molecules (RNA, peptides) on a regular mesh of electric charges at the surface of pyrite that may have
facilitated the primeval polymerization by attracting reactants and arranging them appropriately relative to
each other.[349] The Zn-world theory specifies and differentiates further.[348][350] Hydrothermal fluids rich
in H2 S interacting with cold primordial ocean (or Darwin's "warm little pond") water leads to the
precipitation of metal sulfide particles. Oceanic vent systems and other hydrothermal systems have a zonal
structure reflected in ancient volcanogenic massive sulfide deposits (VMS) of hydrothermal origin. They
reach many kilometers in diameter and date back to the Archean Eon. Most abundant are pyrite (FeS2 ),
chalcopyrite (CuFeS2 ), and sphalerite (ZnS), with additions of galena (PbS) and alabandite (MnS). ZnS
and MnS have a unique ability to store radiation energy, e.g. from UV light. During the relevant time
window of the origins of replicating molecules, the primordial atmospheric pressure was high enough
(>100 bar, about 100 atmospheres) to precipitate near the Earth's surface, and UV irradiation was 10 to 100
times more intense than now; hence the unique photosynthetic properties mediated by ZnS provided just
the right energy conditions to energize the synthesis of informational and metabolic molecules and the
selection of photostable nucleobases.

The Zn-world theory has been further filled out with experimental and theoretical evidence for the ionic
constitution of the interior of the first proto-cells before archaea, bacteria and proto-eukaryotes evolved.
Archibald Macallum noted the resemblance of body fluids such as blood and lymph to seawater;[351]
however, the inorganic composition of all cells differ from that of modern seawater, which led Mulkidjanian
and colleagues to reconstruct the "hatcheries" of the first cells combining geochemical analysis with
phylogenomic scrutiny of the inorganic ion requirements of universal components of modern cells. The
authors conclude that ubiquitous, and by inference primordial, proteins and functional systems show
3−
affinity to and functional requirement for K+, Zn2+, Mn2+, and [PO4 ] . Geochemical reconstruction
shows that the ionic composition conducive to the origin of cells could not have existed in what we today
call marine settings but is compatible with emissions of vapor-dominated zones of what we today call
inland geothermal systems. Under the oxygen depleted, CO2 -dominated primordial atmosphere, the
chemistry of water condensates and exhalations near geothermal fields would resemble the internal milieu
of modern cells. Therefore, the precellular stages of evolution may have taken place in shallow "Darwin
ponds" lined with porous silicate minerals mixed with metal sulfides and enriched in K+, Zn2+, and
phosphorus compounds.[352][353]

Other abiogenesis scenarios


We define a scenario as a set of related concepts pertinent to the origin of life that is or has been
investigated. The concepts related to the Iron-Sulfur world can be considered as a scenario. We consider
some other scenarios that may partially overlap with scenarios discussed above or with each other.

Chemical pathways described by computer

In September 2020, chemists described, for the first time, possible chemical pathways from nonliving
prebiotic chemicals to complex biochemicals that could give rise to living organisms, based on a new
computer program named AllChemy.[354][355]

The hypercycle

In the early 1970s, Manfred Eigen and Peter Schuster examined the transient stages between the molecular
chaos and a self-replicating hypercycle in a prebiotic soup.[356] In a hypercycle, the information storing
system (possibly RNA) produces an enzyme, which catalyzes the formation of another information system,
in sequence until the product of the last aids in the formation of the first information system. Mathematically
treated, hypercycles could create quasispecies, which through natural selection entered into a form of
Darwinian evolution. A boost to hypercycle theory was the discovery of ribozymes capable of catalyzing
their own chemical reactions. The hypercycle theory requires the existence of complex biochemicals, such
as nucleotides, which do not form under the conditions proposed by the Miller–Urey experiment.

Organic pigments in dissipative structures

In his "Thermodynamic Dissipation Theory of the Origin and Evolution of Life",[357][291][293] Karo
Michaelian has taken the insight of Boltzmann and the work of Prigogine to its ultimate consequences
regarding the origin of life. This theory postulates that the hallmark of the origin and evolution of life is the
microscopic dissipative structuring of organic pigments and their proliferation over the entire Earth
surface.[293] Present day life augments the entropy production of Earth in its solar environment by
dissipating ultraviolet and visible photons into heat through organic pigments in water. This heat then
catalyzes a host of secondary dissipative processes such as the water cycle, ocean and wind currents,
hurricanes, etc.[291][294] Michaelian argues that if the thermodynamic function of life today is to produce
entropy through photon dissipation in organic pigments, then this probably was its function at its very
beginnings. It turns out that both RNA and DNA when in water solution are very strong absorbers and
extremely rapid dissipaters of ultraviolet light within the 230–290 nm wavelength (UV-C) region, which is
a part of the Sun's spectrum that could have penetrated the prebiotic atmosphere.[358] In fact, not only RNA
and DNA, but many fundamental molecules of life (those common to all three domains of life) are also
pigments that absorb in the UV-C, and many of these also have a chemical affinity to RNA and DNA.[359]
Nucleic acids may thus have acted as acceptor molecules to the UV-C photon excited antenna pigment
donor molecules by providing an ultrafast channel for dissipation. Michaelian has shown using the
formalism of non-linear irreversible thermodynamics that there would have existed during the Archean a
thermodynamic imperative to the abiogenic UV-C photochemical synthesis and proliferation of these
pigments over the entire Earth surface if they acted as catalysts to augment the dissipation of the solar
photons.[360] By the end of the Archean, with life-induced ozone dissipating UV-C light in the Earth's
upper atmosphere, it would have become ever more improbable for a completely new life to emerge that
did not rely on the complex metabolic pathways already existing since now the free energy in the photons
arriving at Earth's surface would have been insufficient for direct breaking and remaking of covalent bonds.
It has been suggested, however, that such changes in the surface flux of ultraviolet radiation due to
geophysical events affecting the atmosphere could have been what promoted the development of
complexity in life based on existing metabolic pathways, for example during the Cambrian explosion[361]
Some of the most difficult problems concerning the origin of life, such as enzyme-less replication of RNA
and DNA,[362] homochirality of the fundamental molecules,[363] and the origin of information encoding in
RNA and DNA,[364] also find an explanation within the same dissipative thermodynamic framework by
considering the probable existence of a relation between primordial replication and UV-C photon
dissipation. Michaelian suggests that it is erroneous to expect to describe the emergence, proliferation, or
even evolution, of life without overwhelming reference to entropy production through the dissipation of a
generalized thermodynamic potential, in particular, the prevailing solar photon flux.

Protein amyloid

A new origin-of-life theory based on self-replicating beta-sheet structures has been put forward by Maury
in 2009.[365][366] The theory suggest that self-replicating and self-assembling catalytic amyloids were the
first informational polymers in a primitive pre-RNA world. The main arguments for the amyloid hypothesis
is based on the structural stability, autocatalytic and catalytic properties, and evolvability of beta-sheet based
informational systems. Such systems are also error correcting[367] and chiroselective.[368]

Fluctuating salinity: dilute and dry-down

Theories of abiogenesis seldom address the caveat raised by Harold Blum:[369] if the key informational
elements of life – proto-nucleic acid chains – spontaneously form duplex structures, then there is no way to
dissociate them.

Somewhere in this cycle work must be done, which means that free energy must be expended.
If the parts assemble themselves on a template spontaneously, work has to be done to take the
replica off; or, if the replica comes off the template of its own accord, work must be done to
put the parts on in the first place.

The Oparin–Haldane conjecture addresses the formation, but not the dissociation, of nucleic acid polymers
and duplexes. However, nucleic acids are unusual because, in the absence of counterions (low salt) to
neutralize the high charges on opposing phosphate groups, the nucleic acid duplex dissociates into single
chains.[370] Early tides, driven by a close moon, could have generated rapid cycles of dilution (high tide,
low salt) and concentration (dry-down at low tide, high salt) that exclusively promoted the replication of
nucleic acids[370] through a process dubbed tidal chain reaction (TCR).[371] This theory has been criticized
on the grounds that early tides may not have been so rapid,[372] although regression from current values
requires an Earth–Moon juxtaposition at around two Ga, for which there is no evidence, and early tides
may have been approximately every seven hours.[373] Another critique is that only 2–3% of the Earth's
crust may have been exposed above the sea until late in terrestrial evolution.[374]

The TCR (tidal chain reaction) theory has mechanistic advantages over thermal association/dissociation at
deep-sea vents because TCR requires that chain assembly (template-driven polymerization) takes place
during the dry-down phase, when precursors are most concentrated, whereas thermal cycling needs
polymerization to take place during the cold phase, when the rate of chain assembly is lowest and
precursors are likely to be more dilute.

A first protein that condenses substrates during thermal cycling:


thermosynthesis
Emergence of chemiosmotic machinery

Today's bioenergetic process of fermentation is carried


out by either the aforementioned citric acid cycle or the
Acetyl-CoA pathway, both of which have been
connected to the primordial Iron–sulfur world.

In a different approach, the thermosynthesis hypothesis


considers the bioenergetic process of chemiosmosis,
which plays an essential role in cellular respiration and
photosynthesis, more basal than fermentation: the ATP
synthase enzyme, which sustains chemiosmosis, is
Convection cells in fluid placed in a gravity field
proposed as the currently extant enzyme most closely
are selforganizing and enable thermal cycling of
related to the first metabolic process.[375][376] the suspended contents in the fluid such as
protocells containing protoenzymes that work
First life needed an energy source to bring about the
on thermal cycling.
condensation reaction that yielded the peptide bonds of
proteins and the phosphodiester bonds of RNA. In a
generalization and thermal variation of the binding
change mechanism of today's ATP synthase, the "first protein" would have bound substrates (peptides,
phosphate, nucleosides, RNA 'monomers') and condensed them to a reaction product that remained bound
until it was released after a temperature change by a thermal unfolding. The primordial first protein would
therefore have strongly resembled the beta subunits of the ATP synthase alpha/beta subunits of today's F1
moiety in the Fo F1 ATP synthase. Note however that today's enzymes function during isothermal
conditions, whereas the hypothetical first protein worked on and during thermal cycling.

The energy source under the thermosynthesis hypothesis was thermal cycling, the result of suspension of
protocells in a convection current, as is plausible in a volcanic hot spring; the convection accounts for the
self-organization and dissipative structure required in any origin of life model. The still ubiquitous role of
thermal cycling in germination and cell division is considered a relic of primordial thermosynthesis.

By phosphorylating cell membrane lipids, this first protein gave a selective advantage to the lipid protocell
that contained the protein. This protein also synthesized a library of many proteins, of which only a minute
fraction had thermosynthesis capabilities. As proposed by Dyson,[14] it propagated functionally: it made
daughters with similar capabilities, but it did not copy itself. Functioning daughters consisted of different
amino acid sequences.

Whereas the iron–sulfur world identifies a circular pathway as the most simple, the thermosynthesis
hypothesis does not even invoke a pathway: ATP synthase's binding change mechanism resembles a
physical adsorption process that yields free energy,[377] rather than a regular enzyme's mechanism, which
decreases the free energy.

The described first protein may be simple in the sense that is requires only a short sequence of conserved
amino acid residues, a sequent sufficient for the appropriate catalytic cleft. In contrast, it has been claimed
that the emergence of cyclic systems of protein catalysts such as required by fermentation is implausible
because of the length of many required sequences.[378]

Pre-RNA world: The ribose issue and its bypass

It is possible that a different type of nucleic acid, such as peptide nucleic acid, threose nucleic acid or glycol
nucleic acid, was the first to emerge as a self-reproducing molecule, only later replaced by RNA.[379][380]
Larralde et al., say that
the generally accepted prebiotic synthesis of ribose, the formose reaction, yields numerous
sugars without any selectivity.[381]

and they conclude that their

results suggest that the backbone of the first genetic material could not have contained ribose
or other sugars because of their instability.

The ester linkage of ribose and phosphoric acid in RNA is known to be prone to hydrolysis.[382]

Pyrimidine ribonucleosides and their respective nucleotides have been prebiotically synthesized by a
sequence of reactions which by-pass the free sugars, and are assembled in a stepwise fashion by using
nitrogenous or oxygenous chemistries. Sutherland has demonstrated high yielding routes to cytidine and
uridine ribonucleotides built from small 2 and 3 carbon fragments such as glycolaldehyde, glyceraldehyde
or glyceraldehyde-3-phosphate, cyanamide and cyanoacetylene. One of the steps in this sequence allows
the isolation of enantiopure ribose aminooxazoline if the enantiomeric excess of glyceraldehyde is 60% or
greater.[383] This can be viewed as a prebiotic purification step, where the said compound spontaneously
crystallized out from a mixture of the other pentose aminooxazolines. Ribose aminooxazoline can then react
with cyanoacetylene in a mild and highly efficient manner to give the alpha cytidine ribonucleotide.
Photoanomerization with UV light allows for inversion about the 1' anomeric centre to give the correct beta
stereochemistry.[384] In 2009 they showed that the same simple building blocks allow access, via
phosphate controlled nucleobase elaboration, to 2',3'-cyclic pyrimidine nucleotides directly, which are
known to be able to polymerize into RNA. This paper also highlights the possibility for the photo-
sanitization of the pyrimidine-2',3'-cyclic phosphates.[385]

RNA structures

While features of self-organization and self-replication are often considered the hallmark of living systems,
there are many instances of abiotic molecules exhibiting such characteristics under proper conditions. Stan
Palasek suggested based on a theoretical model that self-assembly of ribonucleic acid (RNA) molecules can
occur spontaneously due to physical factors in hydrothermal vents.[386] Virus self-assembly within host
cells has implications for the study of the origin of life,[387] as it lends further credence to the hypothesis
that life could have started as self-assembling organic molecules.[388][389]

Viral origin

Recent evidence for a "virus first" hypothesis, which may support theories of the RNA world, has been
suggested.[390][391] One of the difficulties for the study of the origins of viruses is their high rate of
mutation;[53] this is particularly the case in RNA retroviruses like HIV.[392] A 2015 study compared protein
fold structures across different branches of the tree of life, where researchers can reconstruct the
evolutionary histories of the folds and of the organisms whose genomes code for those folds. They argue
that protein folds are better markers of ancient events as their three-dimensional structures can be
maintained even as the sequences that code for those begin to change.[390] Thus, the viral protein repertoire
retain traces of ancient evolutionary history that can be recovered using advanced bioinformatics
approaches. Those researchers think that "the prolonged pressure of genome and particle size reduction
eventually reduced virocells into modern viruses (identified by the complete loss of cellular makeup),
meanwhile other coexisting cellular lineages diversified into modern cells."[393] The data suggest that
viruses originated from ancient cells that co-existed with the ancestors of modern cells. These ancient cells
likely contained segmented RNA genomes.[390][394]

A computational model (2015) has shown that virus capsids may have originated in the RNA world and
that they served as a means of horizontal transfer between replicator communities since these communities
could not survive if the number of gene parasites increased, with certain genes being responsible for the
formation of these structures and those that favored the survival of self-replicating communities.[395] The
displacement of these ancestral genes between cellular organisms could favor the appearance of new
viruses during evolution.[396] Viruses retain a replication module inherited from the prebiotic stage since it
is absent in cells.[396] So this is evidence that viruses could originate from the RNA world and could also
emerge several times in evolution through genetic escape in cells.[396]

RNA world

A number of hypotheses of formation of RNA have been put forward. As


of 1994, there were difficulties in the explanation of the abiotic synthesis
of the nucleotides cytosine and uracil.[397] Subsequent research has shown
possible routes of synthesis; for example, formamide produces all four
ribonucleotides and other biological molecules when warmed in the
presence of various terrestrial minerals.[220][221] Early cell membranes
could have formed spontaneously from proteinoids, which are protein-like
molecules produced when amino acid solutions are heated while in the
correct concentration of aqueous solution. These are seen to form micro-
spheres which are observed to behave similarly to membrane-enclosed
compartments. Other possible means of producing more complicated
organic molecules include chemical reactions that take place on clay
substrates or on the surface of the mineral pyrite. Jack Szostak

Factors supporting an important role for RNA in early life include its
ability to act both to store information and to catalyze chemical reactions (as a ribozyme); its many
important roles as an intermediate in the expression of and maintenance of the genetic information (in the
form of DNA) in modern organisms; and the ease of chemical synthesis of at least the components of the
RNA molecule under the conditions that approximated the early Earth.[398]

Relatively short RNA molecules have been synthesized, capable of replication.[399] Such replicase RNA,
which functions as both code and catalyst provides its own template upon which copying can occur.
Szostak has shown that certain catalytic RNAs can join smaller RNA sequences together, creating the
potential for self-replication. If these conditions were present, Darwinian natural selection would favour the
proliferation of such autocatalytic sets, to which further functionalities could be added.[400] Such
autocatalytic systems of RNA capable of self-sustained replication have been identified.[401] The RNA
replication systems, which include two ribozymes that catalyze each other's synthesis, showed a doubling
time of the product of about one hour, and were subject to natural selection under the conditions that existed
in the experiment.[402] In evolutionary competition experiments, this led to the emergence of new systems
which replicated more efficiently.[18] This was the first demonstration of evolutionary adaptation occurring
in a molecular genetic system.[402]

Depending on the definition, life started when RNA chains began to self-replicate, initiating the three
mechanisms of Darwinian selection: heritability, variation of type, and differential reproductive output. The
fitness of an RNA replicator (its per capita rate of increase) would likely be a function of its intrinsic
adaptive capacities, determined by its nucleotide sequence, and the availability of resources.[403][404] The
three primary adaptive capacities may have been: (1) replication with moderate fidelity, giving rise to both
heritability while allowing variation of type, (2) resistance to decay, and (3) acquisition of process
resources.[403][404] These capacities would have functioned by means of the folded configurations of the
RNA replicators resulting from their nucleotide sequences.

RNA-DNA world

In the latter half of 2020, evidence, based on a plausibly prebiotic simple compound named
diamidophosphate (DAP), supporting the notion of a RNA-DNA mixture coevolution, has been
presented.[405][406][407][408] The mixture of RNA-DNA sequences, called chimeras, have weak affinity
and form weaker duplex structures.[409] This property is advantageous in an abiotic scenario and these
chimeras have been shown to replicate RNA and DNA – overcoming the "template-product" inhibition
problem, where a pure RNA or pure DNA strand is unable to replicate non-enzymatically because it binds
too strongly to its partners.[410] This behavior of chimeric RNA-DNA sequences could lead to an abiotic
cross-catalytic amplification of RNA and DNA—a key step toward the simultaneous emergence of RNA
and DNA.

Experiments on the origin of life


Both Eigen and Sol Spiegelman demonstrated that evolution, including
replication, variation, and natural selection, can occur in populations of
molecules as well as in organisms.[104] Following on from chemical
evolution came the initiation of biological evolution, which led to the first
cells.[104] No one has yet synthesized a "protocell" using simple
components with the necessary properties of life (the so-called "bottom-up-
approach"). Without such a proof-of-principle, explanations have tended to
focus on chemosynthesis.[411] However, some researchers work in this
field, notably Steen Rasmussen and Szostak, have argued that a "top-down
approach" is more feasible, starting with simple forms of current life.
Spiegelman took advantage of natural selection to synthesize the
J. Craig Venter
Spiegelman Monster, which had a genome with just 218 nucleotide bases,
having deconstructively evolved from a 4500-base bacterial RNA. Eigen
built on Spiegelman's work and produced a similar system further degraded to just 48 or 54 nucleotides—
the minimum required for the binding of the replication enzyme.[412] Craig Venter and others at the J. Craig
Venter Institute engineered existing prokaryotic cells with progressively fewer genes, attempting to discern
at which point the most minimal requirements for life are reached.[413][414][415]

In October 2018, researchers at McMaster University announced the development of a new technology,
called a Planet Simulator, to help study the origin of life on planet Earth and beyond.[416][417][418][419] It
consists of a sophisticated climate chamber to study how the building blocks of life were assembled and
how these prebiotic molecules transitioned into self-replicating RNA molecules.[416]

See also
Anthropic principle – Philosophical premise that all scientific observations presuppose a
universe compatible with the emergence of sentient organisms that make those
observations
Artificial cell
Artificial life – Field of study
Bathybius haeckelii
Entropy and life – Relationship between the thermodynamic concept of entropy and the
evolution of living organisms
Formamide-based prebiotic chemistry
GADV-protein world hypothesis
Hemolithin – Protein claimed to be of extraterrestrial origin
Hypothetical types of biochemistry – Possible alternative biochemicals used by life forms
Mediocrity principle – philosophical concept
Nexus for Exoplanet System Science – Dedicated to the search for life on exoplanets
Noogenesis
Planetary habitability – Extent to which a planet is suitable for life as we know it
Protocell – Lipid globule proposed as a precursor of living cells
Rare Earth hypothesis – Hypothesis that complex extraterrestrial life is improbable and
extremely rare
Shadow biosphere – Hypothetical microbial biosphere of Earth that would use radically
different biochemical and molecular processes from that of currently known life
Tholin – Class of molecules formed by ultraviolet irradiation of organic compounds

References
Footnotes

a. Also occasionally called biopoiesis (Bernal, 1960, p. 30)


b. The reactions are:
Reaction 1: Fayalite + water → magnetite + aqueous silica + hydrogen

3Fe2SiO4 + 2H2O → 2Fe3O4 + 3SiO2 + 2H2

Reaction 2: Forsterite + aqueous silica → serpentine

3Mg2SiO4 + SiO2 + 4H2O → 2Mg3Si2O5(OH)4

Reaction 3: Forsterite + water → serpentine + brucite

2Mg2SiO4 + 3H2O → Mg3Si2O5(OH)4 + Mg(OH)2

Reaction 3 describes the hydration of olivine with water only to yield serpentine and
Mg(OH)2 (brucite). Serpentine is stable at high pH in the presence of brucite like calcium
silicate hydrate, (C-S-H) phases formed along with portlandite (Ca(OH)2) in hardened
Portland cement paste after the hydration of belite (Ca2SiO4), the artificial calcium
equivalent of forsterite. Analogy of reaction 3 with belite hydration in ordinary Portland
cement: Belite + water → C-S-H phase + portlandite

2 Ca2SiO4 + 4 H2O → 3 CaO · 2 SiO2 · 3 H2O + Ca(OH)2

c. The reactions are:

FeS + H2S → FeS2 + 2H+ + 2e−


FeS + H2S + CO2 → FeS2 + HCOOH

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Further reading
"Minerals and the Emergence of Life, pp 135-157 in "Metals, Microbes and Minerals: The
Biogeochemical Side of Life" (2021) pp xiv + 341. Walter de Gruyter, Berlin. Authors Duval,
Simon; Authors Zuchan, Kilian; Baymann, Frauke; Schoepp-Cothenet, Barbara; Branscomb,
Elbert; Russell, Michael, J.; Nitschke, Wolfgang; Editors Kroneck, Peter M.H. and Sosa
Torres, Martha. DOI 10.1515/9783110589771-005 (https://www.degruyter.com/document/doi/
10.1515/9783110589771-005)
Tim Flannery, "In the Soup" (review of Michael Marshall, The Genesis Quest: The Geniuses
and Eccentrics on a Journey to Uncover the Origins of Life on Earth, University of Chicago
Press, 360 pp.), The New York Review of Books, vol. LXVII, no. 19 (3 December 2020),
pp. 37–38.

External links
"Exploring Life's Origins: A Virtual Exhibit" (http://exploringorigins.org/). Exploring Life's
Origins: A Virtual Exhibit. Arlington County, VA: National Science Foundation. Retrieved
2 July 2015.
"The Geochemical Origins of Life by Michael J. Russell & Allan J. Hall" (http://www.gla.ac.u
k/projects/originoflife/). Glasgow, Scotland: University of Glasgow. 13 December 2008.
Retrieved 2 July 2015.
The Origins of Life (https://www.bbc.co.uk/programmes/p004y29f), BBC Radio 4 discussion
with Richard Dawkins, Richard Corfield & Linda Partridge (In Our Time, 23 September
2004)
Minerals and the Origins of Life (https://webcast.stsci.edu/webcast/detail.xhtml?talkid=4006)
(Robert Hazen, NASA) (video, 60m, April 2014).
How life began on Earth (http://www.earthfacts.com/evolution-and-life/howlifebeganearth/)
(Marcia Malory, Earth Facts) (2015)
Stephen Hawking's essay on Life in the Universe (1996) (https://www.hawking.org.uk/in-wor
ds/lectures/life-in-the-universe)
Winston Churchill's essay on Evolution (c.1939/1950s) (https://blogs.scientificamerican.com/
observations/winston-churchills-thoughts-on-evolution/)

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