Abiogenesis

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Abiogenesis

In biology, abiogenesis (from Greek ἀ- a-


'not' + βῐ́ος bios 'life' + γένεσις genesis
'origin') or the origin of life is the natural
process by which life has arisen from non-
living matter, such as simple organic
compounds. The prevailing scientific
hypothesis is that the transition from non-
living to living entities on Earth was not a
single event, but a process of increasing
complexity involving the formation of a
habitable planet, the prebiotic synthesis of
organic molecules, molecular self-
replication, self-assembly, autocatalysis,
and the emergence of cell membranes.
Many proposals have been made for
different stages of the process, but the
transition from non-life to life has never
been observed experimentally.

Stages in the origin of life range from the well-understood, such as


the habitable Earth and the abiotic synthesis of simple molecules, to
the largely unknown, like the derivation of the last universal common
ancestor (LUCA) with its complex molecular functionalities.[1]
The study of abiogenesis aims to
determine how pre-life chemical reactions
gave rise to life under conditions strikingly
different from those on Earth today. It
primarily uses tools from biology and
chemistry, with more recent approaches
attempting a synthesis of many sciences.
Life functions through the specialized
chemistry of carbon and water, and builds
largely upon four key families of
chemicals: lipids for cell membranes,
carbohydrates such as sugars, amino
acids for protein metabolism, and nucleic
acid DNA and RNA for the mechanisms of
heredity. Any successful theory of
abiogenesis must explain the origins and
interactions of these classes of
molecules. 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, although
other self-replicating molecules may have
preceded RNA.

The classic 1952 Miller–Urey experiment


demonstrated that most amino acids, the
chemical constituents of proteins, can be
synthesized from inorganic compounds
under conditions intended to replicate
those of the early Earth. External sources
of energy may have triggered these
reactions, including lightning, radiation,
atmospheric entries of micro-meteorites
and implosion of bubbles in sea and ocean
waves. 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.

A genomics approach has sought to


characterise the last universal common
ancestor (LUCA) of modern organisms by
identifying the genes shared by Archaea
and Bacteria, members of the two major
branches of life (where the Eukaryotes
belong to the archaean branch in the two-
domain system). 355 genes appear to be
common to all life; their nature implies that
the LUCA was anaerobic with the Wood–
Ljungdahl pathway, deriving energy by
chemiosmosis, and maintaining its
hereditary material with DNA, the genetic
code, and ribosomes. Although the LUCA
lived over 4 billion years ago (4 Gya),
researchers do not believe it was the first
form of life. Earlier cells might have had a
leaky membrane and been powered by a
naturally occurring proton gradient near a
deep-sea white smoker hydrothermal vent.
Earth remains the only place in the
universe known to harbor life, and fossil
evidence from the Earth informs most
studies of abiogenesis. The Earth was
formed 4.54 Gya; the earliest undisputed
evidence of life on Earth dates from at
least 3.5 Gya. Fossil micro-organisms
appear to have lived within hydrothermal
vent precipitates dated 3.77 to 4.28 Gya
from Quebec, soon after ocean formation
4.4 Gya during the Hadean.
Overview

NASA's 2015 strategy for astrobiology aimed to solve the


puzzle of the origin of life – how a fully functioning living
system could emerge from non-living components – through
research on the prebiotic origin of life's chemicals, both in
space and on planets, as well as the functioning of early
biomolecules to catalyse reactions and support
inheritance.[2]

Life consists of reproduction with


(heritable) variations.[3] NASA defines life
as "a self-sustaining chemical system
capable of Darwinian [i.e., biological]
evolution."[4] Such a system is complex;
the last universal common ancestor
(LUCA), presumably a single-celled
organism which lived some 4 billion years
ago, already had hundreds of genes
encoded in the DNA genetic code that is
universal today. That in turn implies a suite
of cellular machinery including messenger
RNA, transfer RNA, and ribosomes to
translate the code into proteins. Those
proteins included enzymes to operate its
anaerobic respiration via the Wood–
Ljungdahl metabolic pathway, and a DNA
polymerase to replicate its genetic
material.[5][6]

The challenge for abiogenesis (origin of


life)[7][8][9] researchers is to explain how
such a complex and tightly interlinked
system could develop by evolutionary
steps, as at first sight all its parts are
necessary to enable it to function. For
example, a cell, whether the LUCA or in a
modern organism, copies its DNA with the
DNA polymerase enzyme, which is in turn
produced by translating the DNA
polymerase gene in the DNA. Neither the
enzyme nor the DNA can be produced
without the other.[10] The evolutionary
process could have involved molecular
self-replication, self-assembly such as of
cell membranes, and autocatalysis.[5][6][11]
Nonetheless the transition of non-life to
life has never been observed
experimentally.[12]
The precursors to the development of a
living cell like the LUCA are clear enough, if
disputed in their details: a habitable world
is formed with a supply of minerals and
liquid water. Prebiotic synthesis creates a
range of simple organic compounds,
which are assembled into polymers such
as proteins and RNA. The process after
the LUCA, too, is readily understood:
biological evolution caused the
development of a wide range of species
with varied forms and biochemical
capabilities. The derivation of living things
such as the LUCA from simple
components, however, is far from
understood.[1]
Although Earth remains the only place
where life is known,[13][14] the science of
astrobiology seeks evidence of life on
other planets. The 2015 NASA strategy on
the origin of life aimed to solve the puzzle
by identifying interactions, intermediary
structures and functions, energy sources,
and environmental factors that contributed
to the diversity, selection, and replication
of evolvable macromolecular systems,[2]
and mapping the chemical landscape of
potential primordial informational
polymers. The advent of polymers that
could replicate, store genetic information,
and exhibit properties subject to selection
was, it suggested, most likely a critical
step in the emergence of prebiotic
chemical evolution.[2] Those polymers
derived, in turn, from simple organic
compounds such as nucleobases, amino
acids, and sugars that could have been
formed by reactions in the
environment.[15][8][16][17] A successful
theory of the origin of life must explain
how all these chemicals came into
being.[18]
Pre-1960s conceptual history

The Miller–Urey experiment was a synthesis of small organic molecules in a


mixture of simple gases in a thermal gradient created by heating (right) and
cooling (left) the mixture at the same time, with electrical discharges.

Spontaneous generation

One ancient view of the origin of life, from


Aristotle until the 19th century, is of
spontaneous generation.[19] This theory
held that "lower" animals were generated
by decaying organic substances, and that
life arose by chance.[20][21] This was
questioned from the 17th century, in works
like Thomas Browne's Pseudodoxia
Epidemica.[22][23] In 1665, Robert Hooke
published the first drawings of a
microorganism. In 1676, Antonie van
Leeuwenhoek drew and described
microorganisms, probably protozoa and
bacteria.[24] Van Leeuwenhoek disagreed
with spontaneous generation, and by the
1680s convinced himself, using
experiments ranging from sealed and
open meat incubation and the close study
of insect reproduction, that the theory was
incorrect.[25] In 1668 Francesco Redi
showed that no maggots appeared in
meat when flies were prevented from
laying eggs.[26] By the middle of the 19th
century, spontaneous generation was
considered disproven.[27][28]

Panspermia

Another ancient idea dating back to


Anaxagoras in the 5th century BC is
panspermia,[29] the idea that life exists
throughout the universe, distributed by
meteoroids, asteroids, comets[30] and
planetoids.[31] It does not attempt to
explain how life originated in itself, but
shifts the origin of life on Earth to another
heavenly body. The advantage is that life is
not required to have formed on each
planet it occurs on, but rather in a more
limited set of locations (potentially even a
single location), and then spread about the
galaxy to other star systems via cometary
or meteorite impact.[32]

"A warm little pond": primordial soup

The idea that life originated from non-


living matter in slow stages appeared in
Herbert Spencer's 1864–1867 book
Principles of Biology, and in William Turner
Thiselton-Dyer's 1879 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,
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." Darwin 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."[33][34][35]
Alexander Oparin in 1924 and J. B. S.
Haldane in 1929 proposed that the first
molecules constituting the earliest cells
slowly self-organized from a primordial
soup, and this theory is called the Oparin–
Haldane hypothesis.[36][37] Haldane
suggested that the Earth's prebiotic
oceans consisted of a "hot dilute soup" in
which organic compounds could have
formed.[21][38] J. D. Bernal showed that
such mechanisms could form most of the
necessary molecules for life from
inorganic precursors.[39] In 1967, he
suggested three "stages": the origin of
biological monomers; the origin of
biological polymers; and the evolution
from molecules to cells.[40][41]

Miller–Urey experiment

In 1952, Stanley Miller and Harold Urey


carried out a chemical experiment to
demonstrate how organic molecules could
have formed spontaneously from
inorganic precursors under prebiotic
conditions like those posited by the
Oparin–Haldane hypothesis. It used a
highly reducing (lacking oxygen) mixture
of gases—methane, ammonia, and
hydrogen, as well as water vapor—to form
simple organic monomers such as amino
acids.[42][43] Bernal said of 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."[44] However, current scientific
consensus describes the primitive
atmosphere as weakly reducing or
neutral,[45][46] diminishing the amount and
variety of amino acids that could be
produced. The addition of iron and
carbonate minerals, present in early
oceans, however, produces a diverse array
of amino acids.[45] Later work has focused
on two other potential reducing
environments: outer space and deep-sea
hydrothermal vents.[47][48][49]

Producing a habitable Earth

Evolutionary history

Early universe with first stars

Soon after the Big Bang, which occurred


roughly 14 Gya, the only chemical
elements present in the universe were
hydrogen, helium, and lithium, the three
lightest atoms in the periodic table. These
elements gradually came together to form
stars. These early stars were massive and
short-lived, producing all the heavier
elements through stellar nucleosynthesis.
Carbon, currently the fourth most
abundant chemical element in the
universe (after hydrogen, helium, and
oxygen), was formed mainly in white dwarf
stars, particularly those bigger than twice
the mass of the sun.[50] As these stars
reached the end of their lifecycles, they
ejected these heavier elements, among
them carbon and oxygen, throughout the
universe. These heavier elements allowed
for the formation of new objects, including
rocky planets and other bodies.[51]
According to the nebular hypothesis, the
formation and evolution of the Solar
System began 4.6 Gya with the
gravitational collapse of a small part of a
giant molecular cloud. 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.[52]

Emergence of Earth

The Earth was formed 4.54 Gya.[53][54] The


Hadean Earth (from its formation until 4
Gya) was at first inhospitable to any living
organisms. During its formation, the Earth
lost a significant part of its initial mass,
and consequentially lacked the gravity to
hold molecular hydrogen and the bulk of
the original inert gases.[55] The
atmosphere consisted largely of water
vapor, nitrogen, and carbon dioxide, with
smaller amounts of carbon monoxide,
hydrogen, and sulfur compounds.[56] The
solution of carbon dioxide in water is
thought to have made the seas slightly
acidic, with a pH of about 5.5.[57] The
Hadean atmosphere has been
characterized as a "gigantic, productive
outdoor chemical laboratory,"[58] similar to
volcanic gases today which still support
some abiotic chemistry.[58]
Oceans may have appeared as soon as
200 million years after the Earth formed, in
a near-boiling (100 C) reducing
environment, as the pH of 5.8 rose rapidly
toward neutral.[59] This scenario has found
support from the dating of 4.404 Gya
zircon crystals from metamorphosed
quartzite of Mount Narryer in Western
Australia.[60] Despite the likely increased
volcanism, the Earth may have been a
water world between 4.4 and 4.3 Gya, with
little if any continental crust, a turbulent
atmosphere, and a hydrosphere subject to
intense ultraviolet light from a T Tauri
stage Sun, from cosmic radiation, and
from continued asteroid and comet
impacts.[61]

The Late Heavy Bombardment hypothesis


posits that the Hadean environment
between 4.28[62] and 3.8 Gya was highly
hazardous to 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.[63] Frequent collisions would have
made photosynthesis unviable.[58][64][65][66]
The periods between such devastating
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 have
occurred only between 3.7 and 4.0 Gya.[67]
However, new lunar surveys and samples
have led scientists, including an architect
of the Nice model, to deemphasize the
significance of the Late Heavy
Bombardment.[68]

If life evolved in the ocean at depths of


more than ten meters, it would have been
shielded both from late impacts and the
then high levels of ultraviolet radiation
from the sun. Geothermically heated
oceanic crust could have yielded far more
organic compounds through deep
hydrothermal vents than the Miller–Urey
experiments indicated.[69] The available
energy is maximized at 100–150 °C, the
temperatures at which hyperthermophilic
bacteria and thermoacidophilic archaea
live.[70]

Earliest evidence of life

Life existed on Earth more than 3.5


Gya,[71][72][73] during the Eoarchean when
sufficient crust had solidified following the
molten Hadean.[74][75][76] The earliest
physical evidence of life 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. The
micro-organisms lived within hydrothermal
vent precipitates, soon after the 4.4 Gya
formation of oceans during the Hadean.
The microbes resembled modern
hydrothermal vent bacteria, supporting the
view that abiogenesis began in such an
environment.[62]

Biogenic graphite has been found in 3.7


Gya metasedimentary rocks from
southwestern Greenland[77] and in
microbial mat fossils from 3.49 Gya
Western Australian sandstone.[78]
Evidence of early life in rocks from Akilia
Island, near the Isua supracrustal belt in
southwestern Greenland, dating to 3.7 Gya,
have shown biogenic carbon isotopes.[79]
In other parts of the Isua 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.[80] In the Pilbara region of
Western Australia, compelling evidence of
early life was found in pyrite-bearing
sandstone in a fossilized beach, with
rounded tubular cells that oxidized sulfur
by photosynthesis in the absence of
oxygen.[81][82] Zircons from Western
Australia imply that life existed on Earth at
least 4.1 Gya.[83]

The Pilbara region of Western Australia


contains the Dresser Formation with rocks
3.48 Gya, including layered structures
called stromatolites. Their modern
counterparts are created by
photosynthetic micro-organisms including
cyanobacteria.[84] These lie within
undeformed hydrothermal-sedimentary
strata; their texture indicates a biogenic
origin. Parts of the Dresser formation
preserve hot springs on land, but other
regions seem to have been shallow
seas.[74] A molecular clock analysis
suggests the LUCA emerged prior to the
Late Heavy Bombardment (3.9 Gya).[85]
Stromatolites in the Siyeh Formation, Glacier
National Park, dated 3.5 Gya, placing them
among the earliest life-forms
Modern
stromatolites
in Shark Bay,

Modern stromatolites in Shark Bay, created


by photosynthetic cyanobacteria

Producing molecules:
prebiotic synthesis
All chemical elements except for hydrogen
and helium derive from stellar
nucleosynthesis. The basic chemical
ingredients of life – the carbon-hydrogen
molecule (CH), the carbon-hydrogen
positive ion (CH+) and the carbon ion (C+)
– were produced by ultraviolet light from
stars.[86] Complex molecules, including
organic molecules, form naturally both in
space and on planets.[87] Organic
molecules on the early Earth could have
had either terrestrial origins, with organic
molecule synthesis driven by impact
shocks or by other energy sources, such
as ultraviolet light, redox coupling, or
electrical discharges; or extraterrestrial
origins (pseudo-panspermia), with organic
molecules formed in interstellar dust
clouds raining down on to the planet.[88][89]
Observed extraterrestrial organic
molecules

An organic compound is a chemical


whose molecules contain carbon. Carbon
is abundant in the Sun, stars, comets, and
in the atmospheres of most planets.[90]
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.[87][91][92] Purine and pyrimidine
nucleobases including guanine, adenine,
cytosine, uracil, and thymine have been
found in meteorites. These could have
provided the materials for DNA and RNA to
form on the early Earth.[93] The amino acid
glycine was found in material ejected from
comet Wild 2; it had earlier been detected
in meteorites.[94] Comets are encrusted
with dark material, thought to be a tar-like
organic substance formed from simple
carbon compounds under ionizing
radiation. A rain of material from comets
could have brought such complex organic
molecules to Earth.[95][96][58] 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.[58]
PAH world hypothesis

The Cat's Paw Nebula is inside the Milky Way Galaxy, in


the constellation Scorpius.
Green areas show regions where radiation from hot stars
collided with large molecules and small dust grains
called "polycyclic aromatic hydrocarbons" (PAHs),
causing them to fluoresce. Spitzer Space Telescope,
2018

Polycyclic aromatic hydrocarbons (PAH)


are the most common and abundant
polyatomic molecules in the observable
universe, and are a major store of
carbon.[90][97][98][99] They seem to have
formed shortly after the Big
Bang,[100][98][99] and are associated with
new stars and exoplanets.[90] They are a
likely constituent of Earth's primordial
sea.[100][98][99] PAHs have been detected in
nebulae,[101] and in the interstellar
medium, in comets, and in meteorites.[90]

The PAH world hypothesis posits PAHs as


precursors to the RNA world.[102] A star,
HH 46-IR, resembling the sun early in its
life, is surrounded by a disk of material
which contains molecules including
cyanide compounds, hydrocarbons, and
carbon monoxide. PAHs in the interstellar
medium can be transformed through
hydrogenation, oxygenation, and
hydroxylation to more complex organic
compounds used in living cells.[103]
Nucleobases and Nucleotides

The majority of organic compounds


introduced on Earth by interstellar dust
particles have helped to form complex
molecules, thanks to their peculiar
surface-catalytic activities.[104][105] Studies
of the 12C/13C isotopic ratios of organic
compounds in the Murchison meteorite
suggest that the RNA component uracil
and related molecules, including xanthine,
were formed extraterrestrially.[106] NASA
studies of meteorites suggest that all four
DNA nucleobases (adenine, guanine and
related organic molecules) have been
formed in outer space.[104][107][108] The
cosmic dust permeating the universe
contains complex organics ("amorphous
organic solids with a mixed aromatic–
aliphatic structure") that could be created
rapidly by stars.[109] Glycolaldehyde, a
sugar molecule and RNA precursor, has
been detected in regions of space
including around protostars and on
meteorites.[110][111]

Laboratory synthesis

As early as the 1860s, experiments


demonstrated that biologically relevant
molecules can be produced from
interaction of simple carbon sources with
abundant inorganic catalysts. The
spontaneous formation of complex
polymers from abiotically generated
monomers under the conditions posited
by the "soup" theory is not straightforward.
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.[112] Biology uses essentially
20 amino acids for its coded protein
enzymes, representing a very small subset
of the structurally possible products. Since
life tends to use whatever is available, an
explanation is needed for why the set used
is so small.[113]

Sugars

The Breslow catalytic cycle for formaldehyde


dimerization and C2-C6 sugar formation

Alexander Butlerov showed in 1861 that


the formose reaction created sugars
including tetroses, pentoses, and hexoses
when formaldehyde is heated under basic
conditions with divalent metal ions like
calcium. R. Breslow proposed that the
reaction was autocatalytic in 1959.[114]

Nucleobases

Nucleobases, such as guanine and


adenine, can be synthesized from simple
carbon and nitrogen sources, such as
hydrogen cyanide (HCN) and
ammonia.[115] Formamide produces all
four ribonucleotides when warmed with
terrestrial minerals. Formamide is
ubiquitous in the Universe, produced by
the reaction of water and HCN. It can be
concentrated by the evaporation of
water.[116][117] HCN is poisonous only to
aerobic organisms (eukaryotes and
aerobic bacteria), which did not yet exist. It
can play roles in other chemical processes
such as the synthesis of the amino acid
glycine.[58]

DNA and RNA components including


uracil, cytosine and thymine can be
synthesized under outer space conditions,
using starting chemicals such as
pyrimidine found in meteorites. Pyrimidine
may have been formed in red giant stars or
in interstellar dust and gas clouds.[118] All
four RNA-bases may be synthesized from
formamide in high-energy density events
like extraterrestrial impacts.[119]
Other pathways for synthesizing bases
from inorganic materials have been
reported.[120] Freezing temperatures are
advantageous for the synthesis of purines,
due to the concentrating effect for key
precursors such as hydrogen cyanide.[121]
However, while adenine and guanine
require freezing conditions for synthesis,
cytosine and uracil may require boiling
temperatures.[122] Seven amino acids and
eleven types of nucleobases formed in ice
when ammonia and cyanide were left in a
freezer for 25 years.[123][124] S-triazines
(alternative nucleobases), pyrimidines
including cytosine and uracil, and adenine
can be synthesized by subjecting a urea
solution to freeze-thaw cycles under a
reductive atmosphere, with spark
discharges as an energy source.[125] The
explanation given for the unusual speed of
these reactions at such a low temperature
is eutectic freezing, which crowds
impurities in microscopic pockets of liquid
within the ice, causing the molecules to
collide more often.[126]
Producing suitable vesicles

The three main structures


composed of phospholipids
form spontaneously by self-
assembly in solution: the
liposome (a closed bilayer),
the micelle and the bilayer.

The lipid world theory postulates that the


first self-replicating object was lipid-
like.[127][128] 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. These
bodies may expand by insertion of
additional lipids, and may spontaneously
split into two offspring of similar size and
composition. The main idea is that the
molecular composition of the lipid bodies
is a preliminary to information storage,
and that evolution led to the appearance of
polymers like RNA that store information.
Studies on vesicles from amphiphiles that
might have existed in the prebiotic world
have so far been limited to systems of one
or two types of amphiphiles.[129]

A lipid bilayer membrane could be


composed of a huge number of
combinations of arrangements of
amphiphiles. The best of these would have
favored the constitution of a
hypercycle,[130][131] 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 natural selection.[132]

A protocell is a self-organized, self-


ordered, spherical collection of lipids
proposed as a stepping-stone to the origin
of life.[129] The theory of classical
irreversible thermodynamics treats self-
assembly under a generalized chemical
potential within the framework of
dissipative systems.[133][134][135]

A central question in evolution is how


simple protocells first arose and differed in
reproductive contribution to the following
generation, thus driving the evolution of
life. A functional protocell has (as of 2014)
not yet been achieved in a laboratory
setting.[136][137][138] Self-assembled
vesicles are essential components of
primitive cells.[129] 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
processes from non-living matter.[139]
Irene Chen and Jack W. Szostak suggest
that elementary protocells can give rise to
cellular behaviors including primitive
forms of differential reproduction,
competition, and energy storage.[137]
Competition for membrane molecules
would favor stabilized membranes,
suggesting a selective advantage for the
evolution of cross-linked fatty acids and
even the phospholipids of today.[137] Such
micro-encapsulation would allow for
metabolism within the membrane and the
exchange of small molecules, while
retaining large biomolecules inside. Such a
membrane is needed for a cell to create its
own electrochemical gradient to store
energy by pumping ions across the
membrane.[140][141] Fatty acid vesicles in
conditions relevant to alkaline
hydrothermal vents can be stabilized by
isoprenoids which are synthesized by the
formose reaction, the advantages and
disadvantages of isoprenoids incorporated
within the lipid bilayer in different
microenvironments might have led to the
divergence of the membranes of archaea
and bacteria.[142]
Producing biology

Energy and entropy

Life requires a loss of entropy, or disorder,


when molecules organize themselves into
living matter. The emergence of life and
increased complexity does not contradict
the second law of thermodynamics, which
states that overall entropy never
decreases, since a living organism creates
order in some places (e.g. its living body)
at the expense of an increase of entropy
elsewhere (e.g. heat and waste
production).[143][144][145]
Multiple sources of energy were available
for chemical reactions on the early Earth.
Heat from geothermal processes is a
standard energy source for chemistry.
Other examples include sunlight,
lightning,[58] atmospheric entries of micro-
meteorites,[146] and implosion of bubbles
in sea and ocean waves.[147] This has been
confirmed by experiments[148][149] and
simulations.[150] Unfavorable reactions can
be driven by highly favorable ones, as in
the case of iron-sulfur chemistry. For
example, this was probably important for
carbon fixation.[a] Carbon fixation by
reaction of CO2 with H2S via iron-sulfur
chemistry is favorable, and occurs at
neutral pH and 100 °C. Iron-sulfur
surfaces, which are abundant near
hydrothermal vents, can drive the
production of small amounts of amino
acids and other biomolecules.[58]

Chemiosmosis

ATP synthase uses the


chemiosmotic proton
gradient to power ATP
synthesis through oxidative
phosphorylation.
In 1961, Peter Mitchell proposed
chemiosmosis as a cell's primary system
of energy conversion. The mechanism,
now ubiquitous in living cells, powers
energy conversion in micro-organisms and
in the mitochondria of eukaryotes, making
it a likely candidate for early life.[151][152]
Mitochondria produce adenosine
triphosphate (ATP), the energy currency of
the cell used to drive cellular processes
such as chemical syntheses. The
mechanism of ATP synthesis involves a
closed membrane in which the ATP
synthase enzyme is embedded. The
energy required to release strongly bound
ATP has its origin in protons that move
across the membrane.[153] In modern cells,
those proton movements are caused by
the pumping of ions across the membrane,
maintaining an electrochemical gradient.
In the first organisms, the gradient could
have been provided by the difference in
chemical composition between the flow
from a hydrothermal vent and the
surrounding seawater,[141] or perhaps
meteoric quinones that were conducive to
the development of chemiosmotic energy
across lipid membranes if at a terrestrial
origin.[154]
Chemiosmotic coupling in the membranes of a mitochondrion

The RNA world

The RNA world hypothesis describes an


early Earth with self-replicating and
catalytic RNA but no DNA or proteins.[155]
Many researchers concur that an RNA
world must have preceded the DNA-based
life that now dominates.[156] However,
RNA-based life may not have been the first
to exist.[157][158] Another model echoes
Darwin's "warm little pond" with cycles of
wetting and drying.[159]

RNA is central to the translation process.


Small RNAs can catalyze all the chemical
groups and information transfers required
for life.[158][160] RNA both expresses and
maintains genetic information in modern
organisms; and the chemical components
of RNA are easily synthesized under the
conditions that approximated the early
Earth, which were very different from
those that prevail today. The structure of
the ribosome has been called the
"smoking gun", with a central core of RNA
and no amino acid side chains within 18 Å
of the active site that catalyzes peptide
bond formation.[161][157][162]

The concept of the RNA world was


proposed in 1962 by Alexander Rich,[163]
and the term was coined by Walter Gilbert
in 1986.[158][164] There were initial
difficulties in the explanation of the abiotic
synthesis of the nucleotides cytosine and
uracil.[165] 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.[116][117]
The RNA world hypothesis proposes that undirected polymerisation led to the
emergence of ribozymes, and in turn to an RNA replicase.

RNA replicase can function as both code


and catalyst for further RNA replication,
i.e. it can be autocatalytic. Jack Szostak
has shown that certain catalytic RNAs can
join smaller RNA sequences together,
creating the potential for self-replication.
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
experimental conditions.[166][167][157] If
such conditions were present on early
Earth, then natural selection would favor
the proliferation of such autocatalytic sets,
to which further functionalities could be
added.[168][169][170] Self-assembly of RNA
may occur spontaneously in hydrothermal
vents.[171][172][173] A preliminary form of
tRNA could have assembled into such a
replicator molecule.[174]

Possible precursors to protein synthesis


include the synthesis of short peptide
cofactors or the self-catalysing 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.[175]

Eugene Koonin has argued that "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."[176]

Phylogeny and LUCA

Starting with the work of Carl Woese from


1977, genomics studies have placed the
last universal common ancestor (LUCA) of
all modern life-forms between Bacteria
and a clade formed by Archaea and
Eukaryota in the phylogenetic tree of life. It
lived over 4 Gya.[177][178] A minority of
studies have placed the LUCA in Bacteria,
proposing that Archaea and Eukaryota are
evolutionarily derived from within
Eubacteria;[179] Thomas Cavalier-Smith
suggested in 2006 that the phenotypically
diverse bacterial phylum Chloroflexota
contained the LUCA.[180]
Phylogenetic tree showing the last universal
common ancestor (LUCA) at the root. The
major clades are the Bacteria on one hand,
and the Archaea and Eukaryota on the other.

In 2016, a set of 355 genes likely present


in the LUCA was identified. A total of 6.1
million prokaryotic genes from Bacteria
and Archaea were sequenced, identifying
355 protein clusters from among 286,514
protein clusters that were probably
common to the LUCA. The results suggest
that the LUCA was anaerobic with a
Wood–Ljungdahl (reductive Acetyl-CoA)
pathway, nitrogen- and carbon-fixing,
thermophilic. Its cofactors suggest
dependence upon an environment rich in
hydrogen, carbon dioxide, iron, and
transition metals. Its genetic material was
probably DNA, requiring the 4-nucleotide
genetic code, messenger RNA, transfer
RNA, and ribosomes to translate the code
into proteins such as enzymes. LUCA likely
inhabited an anaerobic hydrothermal vent
setting in a geochemically active
environment. It was evidently already a
complex organism, and must have had
precursors; it was not the first living
thing.[10][181] The physiology of LUCA has
been in dispute.[182][183][184]

LUCA systems and environment included the


Wood–Ljungdahl pathway.[10]
Leslie Orgel argued that early translation
machinery for the genetic code would be
susceptible to error catastrophe. Geoffrey
Hoffmann however showed that such
machinery can be stable in function
against "Orgel's paradox".[185][186][187]
Metabolic reactions that have also been
inferred in LUCA are the incomplete
reverse Krebs cycle, gluconeogenesis, the
pentose phosphate pathway, glycolysis,
reductive amination, and
transamination.[188][189]
Suitable geological
environments

Deep sea hydrothermal vents

The earliest known life forms are putative fossilized


microorganisms, found in white smoker hydrothermal
vent precipitates. They may have lived as early as 4.28
Gya (billion years ago), relatively soon after the formation
of the oceans 4.41 Gya, not long after the formation of
the Earth 4.54 Gya.[62]

Early micro-fossils may have come from a


hot world of gases such as methane,
ammonia, carbon dioxide, and hydrogen
sulfide, toxic to much current life.[190]
Analysis of the tree of life places
thermophilic and hyperthermophilic
bacteria and archaea closest to the root,
suggesting that life may have evolved in a
hot environment.[191] The deep sea or
alkaline hydrothermal vent theory posits
that life began at submarine hydrothermal
vents.[192][193] 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".[194]
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.[192][b]
Early cell powered by external proton gradient near a deep-sea hydrothermal
vent. As long as the membrane (or passive ion channels within it) is
permeable to protons, the mechanism can function without ion pumps.[141]

Russell demonstrated that alkaline vents


created an abiogenic proton motive force
chemiosmotic gradient,[194] ideal for
abiogenesis. 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.[195] 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).[141]

The surfaces of mineral particles inside


deep-ocean hydrothermal vents have
catalytic properties similar to those of
enzymes and can create simple organic
molecules, such as methanol (CH3OH) and
formic, acetic, and pyruvic acids out of the
dissolved CO2 in the water, if driven by an
applied voltage or by reaction with H2 or
H2S.[196][197]
The research reported by Martin in 2016
supports the thesis that life arose at
hydrothermal vents,[198][199] that
spontaneous chemistry in the Earth's crust
driven by rock–water interactions at
disequilibrium thermodynamically
underpinned life's origin[200][201] and that
the founding lineages of the archaea and
bacteria were H2-dependent autotrophs
that used CO2 as their terminal acceptor in
energy metabolism.[202] Martin suggests,
based upon this evidence, that the LUCA
"may have depended heavily on the
geothermal energy of the vent to
survive".[10] Pores at deep sea
hydrothermal vents are suggested to have
been occupied by membrane-bound
compartments which promoted
biochemical reactions.[203][204] Metabolic
intermediates in the Krebs cycle,
gluconeogenesis, amino acid bio-synthetic
pathways, glycolysis, the pentose
phosphate pathway, and including sugars
like ribose, and lipid precursors can occur
non-enzymatically at conditions relevant to
deep-sea alkaline hydrothermal vents.[205]

Hot springs

Mulkidjanian and co-authors think that


marine environments did not provide the
ionic balance and composition universally
found in cells, or the ions required by
essential proteins and ribozymes,
especially with respect to high K+/Na+
ratio, Mn2+, Zn2+ and phosphate
concentrations. They argue that the only
environments that mimic the needed
conditions on Earth are hot springs similar
to ones at Kamchatka.[206] Mineral
deposits in these environments under an
anoxic atmosphere would have suitable
pH (while current pools in an oxygenated
atmosphere would not), contain
precipitates of photocatalytic sulfide
minerals that absorb harmful ultraviolet
radiation, have wet-dry cycles that
concentrate substrate solutions to
concentrations amenable to spontaneous
formation of biopolymers[207][208] created
both by chemical reactions in the
hydrothermal environment, and by
exposure to UV light during transport from
vents to adjacent pools that would
promote the formation of
biomolecules.[209] The hypothesized pre-
biotic environments are similar to
hydrothermal vents, with additional
components that help explain peculiarities
of the LUCA.[206][154]

A phylogenomic and geochemical analysis


of proteins plausibly traced to the LUCA
shows that the ionic composition of its
intracellular fluid is identical to hot
springs. The LUCA likely was dependent
upon synthesized organic matter for its
growth.[206] Experiments show that RNA-
like polymers can be synthesized in wet-
dry cycling and UV light exposure. These
polymers were encapsulated in vesicles
after condensation, which would not
happen in saltwater because of the high
concentrations of ionic solutes.[210]
Potential sources of organics at hot
springs might have been transport by
interplanetary dust particles,
extraterrestrial projectiles, or atmospheric
or geochemical synthesis. Hot springs
could have been abundant at volcanic
landmasses during the Hadean.[154]

Clay

The clay hypothesis was proposed by


Graham Cairns-Smith in 1985.[211][212] It
postulates that complex organic
molecules arose gradually on pre-existing,
non-organic replication surfaces of silicate
crystals in contact with an aqueous
solution. The clay mineral montmorillonite
has been shown to catalyze the
polymerization of RNA in aqueous solution
from nucleotide monomers,[213] and the
formation of membranes from lipids.[214]
In 1998, Hyman Hartman proposed that
"the first organisms were self-replicating
iron-rich clays which fixed carbon dioxide
into oxalic acid and other dicarboxylic
acids. This system of replicating clays and
their metabolic phenotype then evolved
into the sulfide rich region of the hot
spring acquiring the ability to fix nitrogen.
Finally phosphate was incorporated into
the evolving system which allowed the
synthesis of nucleotides and
phospholipids."[215]
Iron–sulfur world

In the 1980s, Wächtershäuser and Karl


Popper postulated the iron–sulfur world
hypothesis for the evolution of pre-biotic
chemical pathways. It traces today's
biochemistry to primordial reactions which
synthesize organic building blocks from
gases.[216][217] Wächtershäuser systems
have a built-in source of energy: iron
sulfides such as pyrite. The energy
released by oxidising these metal sulfides
can support synthesis of organic
molecules. Such systems may have
evolved into autocatalytic sets constituting
self-replicating, metabolically active
entities predating modern life forms.[218]
Experiments with sulfides in an aqueous
environment at 100 °C produced a small
yield of dipeptides (0.4% to 12.4%) and a
smaller yield of tripeptides (0.10%).
However, under the same conditions,
dipeptides were quickly broken down.[219]

Several models postulate a primitive


metabolism, allowing RNA replication to
emerge later. 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.[195]
Concordantly, geochemists Szostak and
Kate Adamala demonstrated that non-
enzymatic RNA replication in primitive
protocells is only possible in the presence
of weak cation chelators like citric acid.
This provides further evidence for the
central role of citric acid in primordial
metabolism.[220] 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).[221][222][218] The physicist
Jeremy England has argued from general
thermodynamic considerations that life
was inevitable.[223] An early version of this
idea was Oparin's 1924 proposal for self-
replicating vesicles. In the 1980s and
1990s came Wächtershäuser's iron–sulfur
world theory and Christian de Duve's
thioester models. More abstract and
theoretical arguments for metabolism
without genes include Freeman Dyson's
mathematical model and Stuart
Kauffman's collectively autocatalytic sets
in the 1980s. Kauffman's work has been
criticized for ignoring the role of energy in
driving biochemical reactions in cells.[224]
The active site of the acetyl-CoA synthase enzyme, part of the acetyl-CoA
pathway, contains nickel-iron-sulfur clusters.

A multistep biochemical pathway like the


Krebs cycle did not just self-organize on
the surface of a mineral; it must have been
preceded by simpler pathways.[225] The
Wood–Ljungdahl pathway is compatible
with self-organization on a metal sulfide
surface. Its key enzyme unit, carbon
monoxide dehydrogenase/acetyl-CoA
synthase, contains mixed nickel-iron-sulfur
clusters in its reaction centers and
catalyzes the formation of acetyl-CoA.
However, prebiotic thiolated and thioester
compounds are thermodynamically and
kinetically unlikely to accumulate in the
presumed prebiotic conditions of
hydrothermal vents.[226] One possibility is
that cysteine and homocysteine may have
reacted with nitriles from the Strecker
reaction, forming catalytic thiol-rich
polypeptides.[227]

It has been suggested that the iron-sulfur


world hypothesis and RNA world
hypothesis are not mutually exclusive as
modern cellular processes do involve both
metabolites and genetic molecules.[228]
Zinc world

Armen Mulkidjanian's zinc world (Zn-


world) hypothesis extends
Wächtershäuser's pyrite
hypothesis.[229][230] The Zn-world theory
proposes that hydrothermal fluids rich in
H2S interacting with cold primordial ocean
(or Darwin's "warm little pond") water
precipitated metal sulfide particles.
Oceanic hydrothermal systems have a
zonal structure reflected in ancient
volcanogenic massive sulfide ore
deposits. They reach many kilometers in
diameter and date back to the Archean.
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 ultraviolet light. When replicating
molecules were originating, the primordial
atmospheric pressure was high enough
(>100 bar) to precipitate near the Earth's
surface, and ultraviolet irradiation was 10
to 100 times more intense than now;
hence the photosynthetic properties
mediated by ZnS provided the right energy
conditions for the synthesis of
informational and metabolic molecules
and the selection of photostable
nucleobases.[229][231]
The Zn-world theory has been filled out
with evidence for the ionic constitution of
the interior of the first protocells. In 1926,
the Canadian biochemist Archibald
Macallum noted the resemblance of body
fluids such as blood and lymph to
seawater;[232] 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 modern cells. The authors conclude
that ubiquitous, and by inference
primordial, proteins and functional
systems show affinity to and functional
requirement for K+, Zn2+, Mn2+, and
3−
[PO4] . Geochemical reconstruction
shows that this ionic composition could
not have existed in the ocean but is
compatible with inland geothermal
systems. In the oxygen-depleted, CO2-
dominated primordial atmosphere, the
chemistry of water condensates near
geothermal fields would resemble the
internal milieu of modern cells. Therefore,
precellular 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.[206][233]
Homochirality

Many biomolecules, such as L-


glutamic acid, are asymmetric, and
occur in living systems in only one of
the two possible forms, in the case of
amino acids the left-handed form.
Prebiotic chemistry would produce
both forms, creating a puzzle for
abiogenesis researchers.[234]

Homochirality is the geometric uniformity


of materials composed of chiral (non-
mirror-symmetric) units. Living organisms
use molecules that have the same chirality
(handedness): with almost no
exceptions,[235] 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 (racemic) mixture of both
forms. 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,[234] and
spontaneous symmetry
breaking.[236][237][238]
Once established, chirality would be
selected for.[239] A small bias
(enantiomeric excess) in the population
can be amplified into a large one by
asymmetric autocatalysis, such as in the
Soai reaction.[240] 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.[241]

Homochirality may have started in outer


space, as on the Murchison meteorite the
amino acid L-alanine is more than twice as
frequent as its D form, and L-glutamic acid
is more than three times as abundant as
its D counterpart.[242][243] Amino acids
from meteorites show a left-handed bias,
whereas sugars show a predominantly
right-handed bias, as found in living
organisms, suggesting an abiogenic origin
of these compounds.[244]

In 2010, an experiment by Robert Root-


Bernstein shows that "two D-RNA-
oligonucleotides having inverse base
sequences (D-CGUA and D-AUGC) and
their corresponding L-RNA-
oligonucleotides (L-CGUA and L-AUGC)
were synthesized and their affinity
determined for Gly and eleven pairs of L-
and D-amino acids". This suggests that
homochirality, including codon
directionality, might have "emerged as a
function of the origin of the genetic
code".[245]

A scenario
The biochemist Nick Lane has proposed a
possible scenario for the origin of life that
integrates much of the available evidence
from biochemistry, geology, phylogeny, and
experimentation:[246]
Iron-Sulphur minerals like Greigite catalyse
the reduction of carbon dioxide in
hydrothermal vents to make Krebs cycle
intermediates.[247]
Protocells in contact with a thin rock barrier
in a hydrothermal vent get a free supply of
energy from the pH gradient.[248]
Protocells in a hydrothermal vent can grow by
adding fatty acids to their membrane, other
organics to their cytoplasm.[249]
Nucleotides in a protocell in a hydrothermal
vent can polymerise into random strings of
RNA. Any that have even slight catalytic
activity will favour the growth and replication
of their protocells, a start to natural
selection.[250]
A protocell away from a hydrothermal vent
must create its own proton-motive force,
such as by splitting hydrogen sulphide.[251]
Ferredoxin catalyses the splitting of hydrogen
sulphide, its earliest repeating amino acid
sequence perhaps coded for by an
incomplete genetic code.[252]
Anoxygenic photosynthesis, using hydrogen
sulphide, ended the need for scarce
hydrogen.[252]
Early heterotrophs used Krebs cycle
respiration; then oxygenic photosynthesis
gave full independence of volcanic
energy.[252]

See also
Autopoiesis
Formamide-based prebiotic chemistry –
Scientific efforts aimed at
reconstructing the beginnings of life
GADV-protein world hypothesis – A
hypothetical stage of abiogenesis
Genetic recombination – Production of
offspring with combinations of traits
that differ from those found in either
parent
Shadow biosphere – Hypothetical
biosphere of Earth

Notes
a. The reactions are:
FeS + H2S → FeS2 + 2H+ + 2e−
FeS + H2S + CO2 → FeS2 + HCOOH

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

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External links
Making headway with the mysteries of
life's origins (https://www.pnas.org/doi/
10.1073/pnas.2105383118) – Adam
Mann (PNAS; 14 April 2021)
Exploring Life's Origins (https://exploring
origins.org/) Archived (https://web.arch
ive.org/web/20230408001731/https://e
xploringorigins.org/) 8 April 2023 at the
Wayback Machine a virtual exhibit at the
Museum of Science (Boston)
How life began on Earth (https://www.ea
rthfacts.com/evolution-and-life/howlifeb
eganearth/) – Marcia Malory (Earth
Facts; 2015)
The Origins of Life (https://www.bbc.co.
uk/programmes/p004y29f) – Richard
Dawkins et al. (BBC Radio; 2004)
Life in the Universe (https://www.hawkin
g.org.uk/in-words/lectures/life-in-the-uni
verse) – Essay by Stephen Hawking
(1996)

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