Origin of Life: PDF Generated At: Tue, 20 Jul 2010 08:43:20 UTC

Origin of Life
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Contents
Articles
Origin of water on Earth 1
Planetary habitability 3
Abiogenesis 18
Miller–Urey experiment 39
Biogenesis 44
Entropy and life 46
Mimivirus 50
Rare Earth hypothesis 54
References
Article Sources and Contributors 65
Image Sources, Licenses and Contributors 67
Article Licenses
License 68
Origin of water on Earth

The question of the origin of water on
Earth, or more accurately put, the question
of why there is clearly more water on the
Earth than on the other planets of the Solar
System, has not been clarified. There are
several acknowledged theories as to how the
world's oceans were formed over the past
4.6 billion years.
Origins Water covers about 70% of the Earth's surface
Some of the most likely contributory factors

to the origin of the Earth's oceans are as follows:
• The cooling of the primordial Earth to the point where the outgassed volatile components were held in an
atmosphere of sufficient pressure for the stabilization and retention of liquid water.
• Comets, trans-Neptunian objects or water-rich meteorites (protoplanets) from the outer reaches of the main
asteroid belt colliding with the Earth may have brought water to the world's oceans. Measurements of the ratio of
the hydrogen isotopes deuterium and protium point to asteroids, since similar percentage impurities in carbon-rich
chondrites were found to oceanic water, whereas previous measurement of the isotopes' concentrations in comets
and trans-Neptunian objects correspond only slightly to water on the earth.
• Biochemically through mineralization and photosynthesis (guttation, transpiration).
• Gradual leakage of water stored in hydrous minerals of the Earth's rocks.
• Photolysis: radiation can break down chemical bonds on the surface.
Water in the development of the Earth

A sizeable quantity of water would have been in the material which formed the Earth.[1] Water molecules would
have escaped Earth's gravity more easily when it was less massive during its formation. Hydrogen and helium are
expected to continually leak from the atmosphere, but the lack of denser noble gases in the modern atmosphere
suggests that something disastrous happened to the early atmosphere.
Part of the young planet is theorized to have been disrupted by the impact which created the Moon, which should
have caused melting of one or two large areas. Present composition does not match complete melting and it is hard to
completely melt and mix huge rock masses.[2] However, a fair fraction of material should have been vaporized by
this impact, creating a rock-vapor atmosphere around the young planet. The rock-vapor would have condensed
within two thousand years, leaving behind hot volatiles which probably resulted in a heavy carbon dioxide
atmosphere with hydrogen and water vapor. Liquid water oceans existed despite the surface temperature of 230°C
because of the atmospheric pressure of the heavy CO2 atmosphere. As cooling continued, subduction and dissolving
in ocean water removed most CO2 from the atmosphere but levels oscillated wildly as new surface and mantle cycles
appeared.[3]
Study of zircons has found that liquid water must have existed as long ago as 4.4 Ga, very soon after the formation
of the Earth.[4] [5] [6] This requires the presence of an atmosphere. The Cool Early Earth theory covers a range from
about 4.4 Ga to 4.0 Ga.
In fact, recent studies of zircons (in the fall of 2008) found in Australian Hadean rock hold minerals that point to the
existence of plate tectonics as early as 4 billion years ago. If this holds true, the previous beliefs about the Hadean
period are far from correct. That is, rather than a hot, molten surface and atmosphere full of carbon dioxide, the
Earth's surface would be very much like it is today. The action of plate tectonics traps vast amounts of carbon
dioxide, thereby eliminating the greenhouse effects and leading to a much cooler surface temperature and the
formation of solid rock, and possibly even life.[7]
Extraterrestrial sources
That the Earth's water originated purely from comets is implausible, as a result of measurements of the isotope ratios
of hydrogen in the three comets Halley, Hyakutake and Hale-Bopp by researchers like David Jewitt, as according to
this research the ratio of deuterium to protium (D/H ratio) of the comets is approximately double that of oceanic
water. What is however unclear is whether these comets are representative of those from the Kuiper Belt. According
to A. Morbidelli [8] the largest part of today's water comes from protoplanets formed in the outer asteroid belt that
plunged towards the Earth, as indicated by the D/H proportions in carbon-rich chondrites. The water in carbon-rich
chondrites point to a similar D/H ratio as oceanic water. Nevertheless, mechanisms have been proposed[9] to suggest
that the D/H-ratio of oceanic water may have increased significantly throughout Earth's history. Such a proposal is
consistent with the possibility that a significant amount of the water on Earth was already present during the planet's
early evolution.
Role of organisms
In the primordial sea's hydrogen sulfide and in the primitive atmosphere present carbon dioxide was used by
sulfide-dependent chemoautotrophic bacteria (prokaryotes) with the supply of light energy for the creation of organic
compounds, whereby water and sulfur resulted:
The greatest proportion of today's water may have been synthesized biochemically through mineralization and
photosynthesis (Calvin cycle).
See also
• Evolution of water on Mars and Earth
Notes
• Jörn Müller, Harald Lesch (2003): Woher kommt das Wasser der Erde? - Urgaswolke oder Meteoriten. Chemie in
unserer Zeit 37(4), pg. 242 – 246, ISSN 0009-2851
• Parts of this article were translated from the original article from the German Wikipedia, on 4/3/06
External links
• Dr. C's Oceans Online website [10] (archived copy [11])
• UniverseToday.com [12]
References
[1] "IngentaConnect Origin of water in the terrestrial planets" (http:/ / www. ingentaconnect. com/ content/ arizona/ maps/ 2005/ 00000040/
00000004/ art00003;jsessionid=7ibpocfkopqql. alice). Ingentaconnect.com. . Retrieved 2009-08-20.
[2] "Solar System Exploration: Science & Technology: Science Features: View Feature" (http:/ / solarsystem. nasa. gov/ scitech/ display.
cfm?ST_ID=446). Solarsystem.nasa.gov. 2004-04-26. . Retrieved 2009-08-20.
[3] N. H. Sleep*,†, K. Zahnle‡, and P. S. Neuhoff§. "Inaugural Article: Initiation of clement surface conditions on the earliest Earth - Sleep et al.
98 (7): 3666 - Proceedings of the National Academy of Sciences" (http:/ / www. pnas. org/ cgi/ content/ full/ 98/ 7/ 3666). Pnas.org. .
Retrieved 2009-08-20.
[4] "ANU - Research School of Earth Sciences - ANU College of Science - Harrison" (http:/ / wwwrses. anu. edu. au/ admin/ index.
php?p=harrison). Ses.anu.edu.au. . Retrieved 2009-08-20.
[5] "ANU - OVC - MEDIA - MEDIA RELEASES - 2005 - NOVEMBER - 181105HARRISONCONTINENTS" (http:/ / info. anu. edu. au/ mac/
Media/ Media_Releases/ _2005/ _November/ _181105harrisoncontinents. asp). Info.anu.edu.au. . Retrieved 2009-08-20.
[6] "A Cool Early Earth" (http:/ / www. geology. wisc. edu/ ~valley/ zircons/ cool_early/ cool_early_home. html). Geology.wisc.edu. . Retrieved
2009-08-20.
[7] Chang, Kenneth (2008-12-02). "A New Picture of the Early Earth" (http:/ / www. nytimes. com/ 2008/ 12/ 02/ science/ 02eart. html?8dpc).
The New York Times. . Retrieved 2010-05-20.
[8] A. Morbidelli et al. Meteoritics & Planetary Science 35, 2000, S. 1309–1329
[9] H. Genda, M. Ikoma, Origin of the Ocean on the Earth: Early Evolution of Water D/H in a Hydrogen-rich Atmosphere. Accessible at http:/ /
arxiv. org/ abs/ 0709. 2025
[10] http:/ / www. oceansonline. com/ ocean_form. htm
[11] http:/ / web. archive. org/ web/ 20050311220523/ http:/ / www. oceansonline. com/ ocean_form. htm
[12] http:/ / www. universetoday. com/ am/ publish/ comets_create_earth_oceans. html?1662004
Planetary habitability
Planetary habitability is the measure of a planet's or a natural
satellite's potential to sustain life. Life may develop directly on a planet
or satellite or be transferred to it from another body, a theoretical
process known as panspermia. As the existence of life beyond Earth is
currently uncertain, planetary habitability is largely an extrapolation of
conditions on Earth and the characteristics of the Sun and solar system
which appear favorable to life's flourishing—in particular those factors
that have sustained complex, multicellular organisms and not just
simpler, unicellular creatures. Research and theory in this regard is a
component of planetary science and the emerging discipline of
astrobiology.
An absolute requirement for life is an energy source, and the notion of Understanding planetary habitability is partly an
extrapolation of the Earth's conditions, as it is the
planetary habitability implies that many other geophysical,
only planet currently known to support life
geochemical, and astrophysical criteria must be met before an
astronomical body can support life. In its astrobiology roadmap, NASA
has defined the principal habitability criteria as "extended regions of liquid water, conditions favorable for the
assembly of complex organic molecules, and energy sources to sustain metabolism."[1]
In determining the habitability potential of a body, studies focus on its bulk composition, orbital properties,
atmosphere, and potential chemical interactions. Stellar characteristics of importance include mass and luminosity,
stable variability, and high metallicity. Rocky, terrestrial-type planets and moons with the potential for Earth-like
chemistry are a primary focus of astrobiological research, although more speculative habitability theories
occasionally examine alternative biochemistries and other types of astronomical bodies.
The idea that planets beyond Earth might host life is an ancient one, though historically it was framed by philosophy
as much as physical science.a The late 20th century saw two breakthroughs in the field. The observation and robotic
spacecraft exploration of other planets and moons within the solar system has provided critical information on
defining habitability criteria and allowed for substantial geophysical comparisons between the Earth and other
bodies. The discovery of extrasolar planets, beginning in the early 1990s[2] [3] and accelerating thereafter, has
provided further information for the study of possible extraterrestrial life. Most importantly, it confirmed that the
Sun is not unique among stars in hosting planets and expanded the habitability research horizon beyond our own
solar system.
Suitable star systems

An understanding of planetary habitability begins with stars. While bodies that are generally Earth-like may be
plentiful, it is just as important that their larger system be agreeable to life. Under the auspices of SETI's Project
Phoenix, scientists Margaret Turnbull and Jill Tarter developed the "HabCat" (or Catalogue of Habitable Stellar
Systems) in 2002. The catalogue was formed by winnowing the nearly 120,000 stars of the larger Hipparcos
Catalogue into a core group of 17,000 "HabStars," and the selection criteria that were used provide a good starting
point for understanding which astrophysical factors are necessary to habitable planets.[4]
Spectral class
The spectral class of a star indicates its photospheric temperature, which (for main-sequence stars) correlates to
overall mass. The appropriate spectral range for "HabStars" is presently considered to be "early F" or "G", to
"mid-K". This corresponds to temperatures of a little more than 7,000 K down to a little more than 4,000 K; the Sun,
a G2 star, is well within these bounds. "Middle-class" stars of this sort have a number of characteristics considered
important to planetary habitability:
• They live at least a few billion years, allowing life a chance to evolve. More luminous main-sequence stars of the
"O," "B," and "A" classes usually live less than a billion years and in exceptional cases less than 10 million.[5] b
• They emit enough high-frequency ultraviolet radiation to trigger important atmospheric dynamics such as ozone
formation, but not so much that ionisation destroys incipient life.[6]
• Liquid water may exist on the surface of planets orbiting them at a distance that does not induce tidal lock (see
next section and 3.2). K Spectrum stars may be able to support life for long periods, far longer than our sun.[7]
This spectral range probably accounts for between 5% and 10% of stars in the local Milky Way galaxy. Whether
fainter late K and M class red dwarf stars are also suitable hosts for habitable planets is perhaps the most important
open question in the entire field of planetary habitability given their ubiquity, see Habitability of red dwarf systems.
Gliese 581 c, a "super-Earth," has been found orbiting in the habitable zone of a red dwarf and may possess liquid
water. Alternately, a greenhouse effect may render it too hot to support life, while its next-nearest neighbor, Gliese
581 d, may in fact be a more likely candidate for habitability.[8]
A stable habitable zone

The habitable zone (HZ) is a theoretical shell surrounding a star in which any planet present would have liquid water
on its surface. After an energy source, liquid water is considered the most important ingredient for life, considering
how integral it is to all life-systems on Earth. This may reflect the bias of a water-dependent species, and if life is
discovered in the absence of water (for example, in a liquid-ammonia solution), the notion of an HZ may have to be
greatly expanded or else discarded altogether as too restricting.c
A "stable" HZ denotes two factors.

First, the range of an HZ should not
vary greatly over time. All stars
increase in luminosity as they age and
a given HZ naturally migrates
outwards, but if this happens too
quickly (for example, with a
super-massive star), planets may only
have a brief window inside the HZ and
a correspondingly weaker chance to
develop life. Calculating an HZ range
and its long-term movement is never
straightforward, given that negative
A range of theoretical habitable zones with stars of different mass (our solar system at
feedback loops such as the carbon centre). Not to scale.
cycle will tend to offset the increases
in luminosity. Assumptions made about atmospheric conditions and geology thus have as great an impact on a
putative HZ range as does Solar evolution; the proposed parameters of the Sun's HZ, for example, have fluctuated
greatly.[9]
Secondly, no large-mass body such as a gas giant should be present in or relatively close to the HZ, thus disrupting
the formation of Earth-like bodies. The mass of the asteroid belt, for example, appears to have been unable to accrete
into a planet due to orbital resonances with Jupiter; if the giant had appeared in the region that is now between the
orbits of Venus and Mars, Earth would almost certainly not have developed its present form. This is somewhat
ameliorated by suggestions that a gas giant inside the HZ might have habitable moons under the right conditions.[10]
The Solar System follows an inner-terrestrial planet, outer-gas giant pattern, but discoveries of extrasolar planets
suggest this may not be common to other stellar systems: numerous Jupiter-sized bodies have been found in close
orbit about their primary, disrupting potential HZs. However, present data for extrasolar planets is likely to be
skewed towards these types (large planets in close orbits) because they are far easier to identify; thus, it remains to
be seen which type of stellar system is the norm, or indeed if there is one.
Low stellar variation

Changes in luminosity are common to all stars, but the severity of such fluctuations covers a broad range. Most stars
are relatively stable, but a significant minority of variable stars often experience sudden and intense increases in
luminosity and consequently the amount of energy radiated toward bodies in orbit. These are considered poor
candidates for hosting life-bearing planets as their unpredictability and energy output changes would negatively
impact organisms. Particularly, living things adapted to a specific temperature range would probably be unable to
survive too great a temperature deviation. Further, upswings in luminosity are generally accompanied by massive
doses of gamma ray and X-ray radiation which might prove lethal. Atmospheres do mitigate such effects (an
absolute increase of 100% in the Sun's luminosity would not necessarily mean a 100% absolute temperature increase
on Earth), but atmosphere retention might not occur on planets orbiting variables, because the high-frequency energy
buffeting these bodies would continually strip them of their protective covering.
The Sun, as in much else, is benign in terms of this danger: the variation between solar max and minimum is roughly
0.1% over its 11-year solar cycle. There is strong (though not undisputed) evidence that even minor changes in the
Sun's luminosity have had significant effects on the Earth's climate well within the historical era; the Little Ice Age
of the mid-second millennium, for instance, may have been caused by a relatively long-term decline in the Sun's
luminosity.[11] Thus, a star does not have to be a true variable for differences in luminosity to affect habitability. Of
known "solar analogs," the one that most closely resembles the Sun is considered to be 18 Scorpii; unfortunately for
the prospects of life existing in its proximity, the only significant difference between the two bodies is the amplitude
of the solar cycle, which appears to be much greater for 18 Scorpii.[12]
High metallicity
While the bulk of material in any star is hydrogen and helium, there is a great variation in the amount of heavier
elements (metals) stars contain. (Note: in this context, "metal" is used in the astronomical context of all elements
heavier than helium, including such elements as carbon, nitrogen, oxygen, phosphorus, and sulfur essential to life as
well as the metals recognized in chemistry). A high proportion of metals in a star correlates to the amount of heavy
material initially available in protoplanetary disks. A low amount of metal significantly decreases the probability that
planets will have formed around that star, under the solar nebula theory of planetary systems formation. Any planets
that did form around a metal-poor star would probably be low in mass, and thus unfavorable for life. Spectroscopic
studies of systems where exoplanets have been found to date confirm the relationship between high metal content
and planet formation: "stars with planets, or at least with planets similar to the ones we are finding today, are clearly
more metal rich than stars without planetary companions."[13] High metallicity also places a requirement for youth
on hab-stars: stars formed early in the universe's history have low metal content and a correspondingly lesser
likelihood of having planetary companions.
Planetary characteristics
The chief assumption about habitable planets is that they are terrestrial.
Such planets, roughly within one order of magnitude of Earth mass, are
primarily composed of silicate rocks and have not accreted the gaseous
outer layers of hydrogen and helium found on gas giants. That life
could evolve in the cloud tops of giant planets has not been decisively
ruled out,d though it is considered unlikely given that they have no
surface and their gravity is enormous.[15] The natural satellites of giant
planets, meanwhile, remain perfectly valid candidates for hosting
life.[14]
The moons of some gas giants could potentially
[14]
In analyzing which environments are likely to support life, a distinction be habitable.
is usually made between simple, unicellular organisms such as bacteria
and archaea and complex metazoans (animals). Unicellularity necessarily precedes multicellularity in any
hypothetical tree of life and where single-celled organisms do emerge there is no assurance that this will lead to
greater complexity.e The planetary characteristics listed below are considered crucial for life generally, but in every
case habitability impediments should be considered greater for multicellular organisms such as plants and animals
versus unicellular life.
Mass
Low-mass planets are poor candidates for life for two reasons. First,
their lesser gravity makes atmosphere retention difficult. Constituent
molecules are more likely to reach escape velocity and be lost to space
when buffeted by solar wind or stirred by collision. Planets without a
thick atmosphere lack the matter necessary for primal biochemistry,
have little insulation and poor heat transfer across their surfaces (for
example, Mars, with its thin atmosphere, is colder than the Earth would
be if it were at a similar distance from the sun), and provide less
protection against meteoroids and high-frequency radiation. Further,
where an atmosphere is less than 0.006 Earth atmospheres, water
cannot exist in liquid form as the required atmospheric pressure, 4.56
mm Hg (608 Pa) (0.18 inch Hg), does not occur. The temperature Mars, with its thin atmosphere, is colder than the
range at which water is liquid is smaller at low pressures generally. Earth would be, if it were at a similar distance
from the Sun
Secondly, smaller planets have smaller diameters and thus higher
surface-to-volume ratios than their larger cousins. Such bodies tend to lose the energy left over from their formation
quickly and end up geologically dead, lacking the volcanoes, earthquakes and tectonic activity which supply the
surface with life-sustaining material and the atmosphere with temperature moderators like carbon dioxide. Plate
tectonics appear particularly crucial, at least on Earth: not only does the process recycle important chemicals and
minerals, it also fosters bio-diversity through continent creation and increased environmental complexity and helps
create the convective cells necessary to generate Earth's magnetic field.[16]
"Low mass" is partly a relative label; the Earth is considered low mass when compared to the Solar System's gas
giants, but it is the largest, by diameter and mass, and densest of all terrestrial bodies.f It is large enough to retain an
atmosphere through gravity alone and large enough that its molten core remains a heat engine, driving the diverse
geology of the surface (the decay of radioactive elements within a planet's core is the other significant component of
planetary heating). Mars, by contrast, is nearly (or perhaps totally) geologically dead and has lost much of its
atmosphere.[17] Thus, it would be fair to infer that the lower mass limit for habitability lies somewhere between that
of Mars and Earth or Venus; 0.3 Earth masses has been offered as a rough dividing line for habitable planets.[18]
However, a 2008 study by the Harvard-Smithsonian Center for Astrophysics suggests that the dividing line may be
higher. Earth may in fact lie on the lower boundary of habitability, since if it were any smaller, plate tectonics would
be impossible. Venus, which has 85 percent Earth's mass, shows no signs of tectonic activity. Conversely,
"super-Earths", terrestrial planets with higher masses than Earth, would have higher levels of plate tectonics and thus
be firmly placed in the habitable range.[19] Exceptional circumstances do offer exceptional cases: Jupiter's moon Io
(which is smaller than any of the terrestrial planets) is volcanically dynamic because of the gravitational stresses
induced by its orbit, and its neighbor Europa may have a liquid ocean underneath a frozen shell also due to power
generated from orbiting a gas giant. Saturn's Titan, meanwhile, has an outside chance of harbouring life, as it has
retained a thick atmosphere and bio-chemical reactions are possible in the liquid methane on its surface. These
satellites are exceptions, but they prove that mass as a habitability criterion cannot be considered definitive.
Finally, a larger planet is likely to have a large iron core. This allows for a magnetic field to protect the planet from
stellar wind and cosmic radiation, which otherwise would tend to strip away planetary atmosphere and to bombard
living things with ionized particles. Mass is not the only criterion for producing a magnetic field—as the planet must
also rotate fast enough to produce a dynamo effect within its core[20] —but it is a significant component of the
process.
Orbit and rotation

As with other criteria, stability is the critical consideration in determining the effect of orbital and rotational
characteristics on planetary habitability. Orbital eccentricity is the difference between a planet's farthest and closest
approach to its parent star divided by the sum of said distances. It is a ratio describing the shape of the elliptical
orbit. The greater the eccentricity the greater the temperature fluctuation on a planet's surface. Although they are
adaptive, living organisms can only stand so much variation, particularly if the fluctuations overlap both the freezing
point and boiling point of the planet's main biotic solvent (e.g., water on Earth). If, for example, Earth's oceans were
alternately boiling and freezing solid, it is difficult to imagine life as we know it having evolved. The more complex
the organism, the greater the temperature sensitivity.[21] The Earth's orbit is almost wholly circular, with an
eccentricity of less than 0.02; other planets in our solar system (with the exception of Mercury) have eccentricities
that are similarly benign.
Data collected on the orbital eccentricities of extrasolar planets has surprised most researchers: 90% have an orbital
eccentricity greater than that found within the solar system, and the average is fully 0.25.[22]
A planet's movement around its rotational axis must also meet certain criteria if life is to have the opportunity to
evolve. A first assumption is that the planet should have moderate seasons. If there is little or no axial tilt (or
obliquity) relative to the perpendicular of the ecliptic, seasons will not occur and a main stimulant to biospheric
dynamism will disappear. The planet would also be colder than it would be with a significant tilt: when the greatest
intensity of radiation is always within a few degrees of the equator, warm weather cannot move poleward and a
planet's climate becomes dominated by colder polar weather systems.
If a planet is radically tilted, meanwhile, seasons will be extreme and make it more difficult for a biosphere to
achieve homeostasis. Although during the Quaternary higher axial tilt of the Earth coincides with reduced polar ice,
warmer temperatures and less seasonal variation, scientists do not know whether this trend would continue
indefinitely with further increases in axial tilt (see Snowball Earth).
The exact effects of these changes can only be computer modelled at present, and studies have shown that even
extreme tilts of up to 85 degrees do not absolutely preclude life "provided it does not occupy continental surfaces
plagued seasonally by the highest temperature."[23] Not only the mean axial tilt, but also its variation over time must
be considered. The Earth's tilt varies between 21.5 and 24.5 degrees over 41,000 years. A more drastic variation, or a
much shorter periodicity, would induce climatic effects such as variations in seasonal severity.
Other orbital considerations include:
• The planet should rotate relatively quickly so that the day-night cycle is not overlong. If a day takes years, the
temperature differential between the day and night side will be pronounced, and problems similar to those noted
with extreme orbital eccentricity will come to the fore.
• Change in the direction of the axis rotation (precession) should not be pronounced. In itself, precession need not
affect habitability as it changes the direction of the tilt, not its degree. However, precession tends to accentuate
variations caused by other orbital deviations; see Milankovitch cycles. Precession on Earth occurs over a 26,000
year cycle.
The Earth's Moon appears to play a crucial role in moderating the Earth's climate by stabilising the axial tilt. It has
been suggested that a chaotic tilt may be a "deal-breaker" in terms of habitability— i.e. a satellite the size of the
moon is not only helpful but required to produce stability.[24] This position remains controversial.g
Geochemistry
It is generally assumed that any extraterrestrial life that might exist will be based on the same fundamental
biochemistry as found on Earth, as the four elements most vital for life, carbon, hydrogen, oxygen, and nitrogen, are
also the most common chemically reactive elements in the universe. Indeed, simple biogenic compounds, such as
amino acids, have been found in meteorites and in the interstellar medium.[25] These four elements together comprise
over 96% of Earth's collective biomass. Carbon has an unparalleled ability to bond with itself and to form a massive
array of intricate and varied structures, making it an ideal material for the complex mechanisms that form living
cells. Hydrogen and oxygen, in the form of water, compose the solvent in which biological processes take place and
in which the first reactions occurred that led to life's emergence. The energy released in the formation of powerful
covalent bonds between carbon and oxygen, available by oxidizing organic compounds, is the fuel of all complex
life-forms. These four elements together make up amino acids, which in turn are the building blocks of proteins, the
substance of living tissue. In addition, neither sulfur, required for the building of proteins, nor phosphorus, needed
for the formation of DNA, RNA, and the adenosine phosphates essential to metabolism, are rare.
Relative abundance in space does not always mirror differentiated abundance within planets; of the four life
elements, for instance, only oxygen is present in any abundance in the Earth's crust.[26] This can be partly explained
by the fact that many of these elements, such as hydrogen and nitrogen, along with their simplest and most common
compounds, such as carbon dioxide, carbon monoxide, methane, ammonia, and water, are gaseous at warm
temperatures. In the hot region close to the Sun, these volatile compounds could not have played a significant role in
the planets' geological formation. Instead, they were trapped as gases underneath the newly formed crusts, which
were largely made of rocky, involatile compounds such as silica (a compound of silicon and oxygen, accounting for
oxygen's relative abundance). Outgassing of volatile compounds through the first volcanoes would have contributed
to the formation of the planets' atmospheres. The Miller-Urey experiment showed that, with the application of
energy, amino acids can form from the synthesis of the simple compounds within a primordial atmosphere.[27]
Even so, volcanic outgassing could not have accounted for the amount of water in Earth's oceans.[28] The vast
majority of the water —and arguably carbon— necessary for life must have come from the outer solar system, away
from the Sun's heat, where it could remain solid. Comets impacting with the Earth in the Solar system's early years
would have deposited vast amounts of water, along with the other volatile compounds life requires (including amino
acids) onto the early Earth, providing a kick-start to the origin of life.
Thus, while there is reason to suspect that the four "life elements" ought to be readily available elsewhere, a
habitable system probably also requires a supply of long-term orbiting bodies to seed inner planets. Without comets
there is a possibility that life as we know it would not exist on Earth.
Microenvironments and extremophiles

One important qualification to habitability criteria is that only a tiny
portion of a planet is required to support life. Astrobiologists often
concern themselves with "micro-environments," noting that "we lack a
fundamental understanding of how evolutionary forces, such as
mutation, selection, and genetic drift, operate in micro-organisms that
act on and respond to changing micro-environments."[29]
Extremophiles are Earth organisms that live in niche environments
under severe conditions generally considered inimical to life. Usually
(although not always) unicellular, extremophiles include acutely
alkaliphilic and acidophilic organisms and others that can survive
water temperatures above 100 °C in hydrothermal vents.
The discovery of life in extreme conditions has complicated definitions

of habitability, but also generated much excitement amongst
researchers in greatly broadening the known range of conditions under
which life can persist. For example, a planet that might otherwise be
unable to support an atmosphere given the solar conditions in its
vicinity, might be able to do so within a deep shadowed rift or volcanic The Atacama Desert provides an analog to Mars
[30] and an ideal environment to study the boundary
cave. Similarly, craterous terrain might offer a refuge for primitive
between sterility and habitability.
life. The Lawn Hill crater has been studied as an astrobiological
analog, with researchers suggesting rapid sediment infill created a
protected microenvironment for microbial organisms; similar conditions may have occurred over the geological
history of Mars.[31]
Earth environments that cannot support life are still instructive to astrobiologists in defining the limits of what
organisms can endure. The heart of the Atacama desert, generally considered the driest place on Earth, appears
unable to support life, but it has been subject to study by NASA for that reason: it provides a Mars analog and the
moisture gradients along its edges are ideal for studying the boundary between sterility and habitability.[32] The
Atacama was the subject of study in 2003 that partly replicated experiments from the Viking landings on Mars in the
1970s; no DNA could be recovered from two soil samples, and incubation experiments were also negative for
biosignatures.[33]
Alternative star systems

In determining the feasibility of extraterrestrial life, astronomers had long focused their attention on stars like our
own Sun. However, they have begun to explore the possibility that life might form in systems very unlike our own.
Binary systems
Typical estimates often suggest that 50% or more of all stellar systems are binary systems. This may be partly
sample bias, as massive and bright stars tend to be in binaries and these are most easily observed and catalogued; a
more precise analysis has suggested that more common, fainter, stars are usually singular and that up to two thirds of
all stellar systems are therefore solitary.[34]
The separation between stars in a binary may range from less than one astronomical unit (AU, the Earth-Sun
distance) to several hundred. In latter instances, the gravitational effects will be negligible on a planet orbiting an
otherwise suitable star and habitability potential will not be disrupted unless the orbit is highly eccentric (see
Nemesis, for example). However, where the separation is significantly less, a stable orbit may be impossible. If a
planet’s distance to its primary exceeds about one fifth of the closest approach of the other star, orbital stability is not
guaranteed.[35] Whether planets might form in binaries at all had long been unclear, given that gravitational forces
might interfere with planet formation. Theoretical work by Alan Boss at the Carnegie Institution has shown that gas
giants can form around stars in binary systems much as they do around solitary stars.[36]
One study of Alpha Centauri, the nearest star system to the Sun, suggested that binaries need not be discounted in the
search for habitable planets. Centauri A and B have an 11 AU distance at closest approach (23 AU mean), and both
should have stable habitable zones. A study of long-term orbital stability for simulated planets within the system
shows that planets within approximately three AU of either star may remain stable (i.e. the semi-major axis deviating
by less than 5%). The HZ for Centauri A is conservatively estimated at 1.2 to 1.3 AU and Centauri B at 0.73 to
0.74—well within the stable region in both cases.[37]
Red dwarf systems

Determining the habitability of red dwarf
stars could help determine how common life
in the universe might be, as red dwarfs make
up between 70 to 90% of all the stars in the
galaxy. Brown dwarfs are probably more
numerous than red dwarfs. However, they
are not generally classified as stars, and
could never support life as we understand it,
since what little heat they emit quickly
disappears.
Astronomers for many years ruled out red

dwarfs as potential abodes for life. Their
small size (from 0.1 to 0.6 solar masses) Relative star sizes and photospheric temperatures. Any planet around a red dwarf
means that their nuclear reactions proceed such as the one shown here would have to huddle close to achieve Earth-like
exceptionally slowly, and they emit very temperatures, probably inducing tidal lock. See Aurelia.
little light (from 3% of that produced by the

Sun to as little as 0.01%). Any planet in orbit around a red dwarf would have to huddle very close to its parent star to
attain Earth-like surface temperatures; from 0.3 AU (just inside the orbit of Mercury) for a star like Lacaille 8760, to
as little as 0.032 AU for a star like Proxima Centauri[38] (such a world would have a year lasting just 6.3 days). At
those distances, the star's gravity would cause tidal lock. One side of the planet would eternally face the star, while
the other would always face away from it. The only way potential life could avoid either an inferno or a deep freeze
would be if the planet had an atmosphere thick enough to transfer the star's heat from the day side to the night side. It
was long assumed that such a thick atmosphere would prevent sunlight from reaching the surface in the first place,
preventing photosynthesis.
This pessimism has been tempered by research. Studies by Robert Haberle and Manoj Joshi of NASA's Ames
Research Center in California have shown that a planet's atmosphere (assuming it included greenhouse gases CO2
and H2O) need only be 100 mbs, or 10% of Earth's atmosphere, for the star's heat to be effectively carried to the
night side.[39] This is well within the levels required for photosynthesis, though water would still remain frozen on
the dark side in some of their models. Martin Heath of Greenwich Community College, has shown that seawater,
too, could be effectively circulated without freezing solid if the ocean basins were deep enough to allow free flow
beneath the night side's ice cap. Further research—including a consideration of the amount of photosynthetically
active radiation—suggested that tidally locked planets in red dwarf systems might at least be habitable for higher
plants.[40]
Size is not the only factor in making red dwarfs potentially unsuitable for life, however. On a red dwarf planet,
photosynthesis on the night side would be impossible, since it would never see the sun. On the day side, because the
sun does not rise or set, areas in the shadows of mountains would remain so forever. Photosynthesis as we
understand it would be complicated by the fact that a red dwarf produces most of its radiation in the infrared, and on
the Earth the process depends on visible light. There are potential positives to this scenario. Numerous terrestrial
ecosystems rely on chemosynthesis rather than photosynthesis, for instance, which would be possible in a red dwarf
system. A static primary star position removes the need for plants to steer leaves toward the sun, deal with changing
shade/sun patterns, or change from photosynthesis to stored energy during night. Because of the lack of a day-night
cycle, including the weak light of morning and evening, far more energy would be available at a given radiation
level.
Red dwarfs are far more variable and violent than their more stable, larger cousins. Often they are covered in
starspots that can dim their emitted light by up to 40% for months at a time, while at other times they emit gigantic
flares that can double their brightness in a matter of minutes.[41] Such variation would be very damaging for life, as it
would not only destroy any complex organic molecules that could possibly form biological precursors, but also
because it would blow off sizeable portions of the planet's atmosphere. For a planet around a red dwarf star to
support life, it would require a rapidly rotating magnetic field to protect it from the flares. However, a tidally locked
planet rotates only very slowly, and so cannot produce a geodynamo at its core. However, the violent flaring period
of a red dwarf's life cycle is estimated to only last roughly the first 1.2 billion years of its existence. If a planet forms
far away from a red dwarf so as to avoid tidal locking, and then migrates into the star's habitable zone after this
turbulent initial period, it is possible that life may have a chance to develop.[42]
There is, however, one major advantage that red dwarfs have over other stars as abodes for life: they live a long time.
It took 4.5 billion years before humanity appeared on Earth, and life as we know it will see suitable conditions for at
most 1 billion years more.[43] Red dwarfs, by contrast, could live for trillions of years because their nuclear reactions
are far slower than those of larger stars, meaning that life would have longer to evolve and survive. Further, while
the odds of finding a planet in the habitable zone around any specific red dwarf are slim, the total amount of
habitable zone around all red dwarfs combined is equal to the total amount around Sun-like stars given their
ubiquity.[44]
The galactic neighborhood

Along with the characteristics of planets and their star systems, the wider galactic environment may also impact
habitability. Scientists considered the possibility that particular areas of galaxies (galactic habitable zones) are better
suited to life than others; the solar system in which we live, in the Orion Spur, on the Milky Way galaxy's edge is
considered to be in a life-favorable spot:[45]
• It is not in a globular cluster where immense star densities are inimical to life, given excessive radiation and
gravitational disturbance. Globular clusters are also primarily composed of older, probably metal-poor, stars.
• It is not near an active gamma ray source.
• It is not near the galactic center where once again star densities increase the likelihood of ionizing radiation (e.g.,
from magnetars and supernovae). A supermassive black hole is also believed to lie at the middle of the galaxy
which might prove a danger to any nearby bodies.
• The circular orbit of the Sun around the galactic centre keeps it out of the way of the galaxy's spiral arms where
once more intense radiation and gravitation may lead to disruption.[46]
Thus, relative loneliness is ultimately what a life-bearing system needs. If the Sun were crowded amongst other
systems, the chance of being fatally close to dangerous radiation sources would increase significantly. Further, close
neighbours might disrupt the stability of various orbiting bodies such as Oort cloud and Kuiper Belt objects, which
can bring catastrophe if knocked into the inner solar system.
While stellar crowding proves disadvantageous to habitability, so too does extreme isolation. A star as metal-rich as
the Sun would probably not have formed in the very outermost regions of the Milky Way given a decline in the
relative abundance of metals and a general lack of star formation. Thus, a "suburban" location, such as our Solar
System enjoys, is preferable to a Galaxy's center or farthest reaches.[47]
Other considerations
Alternative biochemistries
While most investigations of extraterrestrial life start with the assumption that advanced life-forms must have similar
requirements for life as on Earth, the hypothesis of other types of biochemistry suggests the possibility of lifeforms
evolving around a different metabolic mechanism. In Evolving the Alien, biologist Jack Cohen and mathematician
Ian Stewart argue astrobiology, based on the Rare Earth hypothesis, is restrictive and unimaginative. They suggest
that Earth-like planets may be very rare, but non-carbon-based complex life could possibly emerge in other
environments. The most frequently mentioned alternative to carbon is silicon-based life, while ammonia is
sometimes suggested as an alternative solvent to water.
More speculative ideas have focused on bodies altogether different than Earth-like planets. Astronomer Frank Drake,
a well-known proponent of the search for extraterrestrial life, imagined life on a neutron star: submicroscopic
"nuclear molecules" combining to form creatures with a life cycle millions of times quicker than Earth life.[48] Called
"imaginative and tongue-in-cheek," the idea gave rise to science fiction depictions.[49] Carl Sagan, another optimist
with regards to extraterrestrial life, considered the possibility of organisms that are always airborne within the high
atmosphere of Jupiter in a 1976 paper.[50] [51] Cohen and Stewart also envisioned life in both a solar environment
and in the atmosphere of a gas giant.
"Good Jupiters"
"Good Jupiters" are gas giant planets, like the solar system's Jupiter, that orbit their stars in circular orbits far enough
away from the habitable zone to not disturb it but close enough to "protect" terrestrial planets in closer orbit in two
critical ways. First, they help to stabilize the orbits, and thereby the climates, of the inner planets. Second, they keep
the inner solar system relatively free of comets and asteroids that could cause devastating impacts.[52] Jupiter orbits
the Sun at about five times the distance between the Earth and the Sun. This is the rough distance we should expect
to find good Jupiters elsewhere. Jupiter's "caretaker" role was dramatically illustrated in 1994 when Comet
Shoemaker-Levy 9 impacted the giant; had Jovian gravity not captured the comet, it may well have entered the inner
solar system.
Early in the Solar System's history, Jupiter played a somewhat contrary role: it increased the eccentricity of asteroid
belt orbits and enabled many to cross Earth's orbit and supply the planet with important volatiles. Before Earth
reached half its present mass, icy bodies from the Jupiter–Saturn region and small bodies from the primordial
asteroid belt supplied water to the Earth due to the gravitational scattering of Jupiter and, to a lesser extent,
Saturn.[53] Thus, while the gas giants are now helpful protectors, they were once suppliers of critical habitability
material.
In contrast, Jupiter-sized bodies that orbit too close to the habitable zone but not in it (as in 47 Ursae Majoris), or
have a highly elliptical orbit that crosses the habitable zone (like 16 Cygni B) make it very difficult for an Earthlike
planet to exist in the system. See the discussion of a stable habitable zone above.
Life's impact on habitability

A supplement to the factors that support life's emergence is the notion that life itself, once formed, becomes a
habitability factor in its own right. An important Earth example was the production of oxygen by ancient
cyanobacteria, and eventually photosynthesizing plants, leading to a radical change in the composition of Earth’s
atmosphere. This oxygen would prove fundamental to the respiration of later animal species.
This interaction between life and subsequent habitability has been explored in various ways. The Gaia hypothesis, a
class of scientific models of the geo-biosphere pioneered by Sir James Lovelock in 1975, argues that life as a whole
fosters and maintains suitable conditions for itself by helping to create a planetary environment suitable for its
continuity; at its most dramatic, Gaia suggests that planetary systems behave similarly to a kind of organism. The
most successful life forms change the composition of the air, water, and soil in ways that make their continued
existence more certain—a controversial extension of the accepted laws of ecology.
Similarly, David Grinspoon has suggested a "Living Worlds hypothesis" in which our understanding of what
constitutes habitability cannot be separated from life already extant on a planet. Planets that are geologically and
meteorologically alive are much more likely to be biologically alive as well and "a planet and its life will
co-evolve."[54]
In their 2004 book The Privileged Planet, astronomer Guillermo Gonzalez and philosopher Jay Richards explore the
possible link between the habitability of a planet and its suitability for observing the rest of the universe. The book
was criticized as an example of intelligent design and for its lack of scientific credibility.[55]
See also
• Alien Planet
• Class M planet
• Darwin mission
• Definition of planet
• Extraterrestrial liquid water
• Goldilocks planet
• Neocatastrophism
• Space colonization
• Terraforming
• Terrestrial Planet Finder
Notes
• Note a: This article is an analysis of planetary habitability from the perspective of contemporary physical science.
A historical viewpoint on the possibility of habitable planets can be found at Beliefs in extraterrestrial life and
Cosmic pluralism. For a discussion of the probability of alien life see the Drake Equation and Fermi Paradox.
Habitable planets are also a staple of fiction; see Planets in science fiction.
• Note b: Life appears to have emerged on Earth approximately 500 million years after the planet’s formation. "A"
class stars (which shine for between 600 million and 1.2 billion years) and a small fraction of "B" class stars
(which shine 10+ million to 600 million) fall within this window. At least theoretically life could emerge in such
systems but it would almost certainly not reach a sophisticated level given these time-frames and the fact that
increases in luminosity would occur quite rapidly. Life around "O" class stars is exceptionally unlikely, as they
shine for less than ten million years.
• Note c: That Europa and to a lesser extent Titan (respectively, 3.5 and 8 astronomical units outside our Sun’s
putative habitable zone) are considered prime extraterrestrial possibilities underscores the problematic nature of
the HZ criterion. In secondary and tertiary descriptions of habitability it is often stated that habitable planets must
be within the HZ—this remains to be proven.

• Note d: In Evolving the Alien, Jack Cohen and Ian Stewart evaluate plausible scenarios in which life might form
in the cloud-tops of Jovian planets. Similarly, Carl Sagan suggested that the clouds of Jupiter might host life.[50]
.[51]
• Note e: There is an emerging consensus that single-celled micro-organisms may in fact be common in the
universe, especially since Earth’s extremophiles flourish in environments that were once considered hostile to life.
The potential occurrence of complex multi-celled life remains much more controversial. In their work Rare
Earth: Why Complex Life Is Uncommon in the Universe, Peter Ward and Donald Brownlee argue that microbial
life is probably widespread while complex life is very rare and perhaps even unique to Earth. Current knowledge
of Earth’s history partly buttresses this theory: multi-celled organisms are believed to have emerged at the time of
the Cambrian explosion close to 600 mya but more than 3 billion years after life itself appeared. That Earth life
remained unicellular for so long underscores that the decisive step toward complex organisms need not
necessarily occur.
• Note f: There is a "mass-gap" in our solar system between Earth and the two smallest gas giants, Uranus and
Neptune, which are 13 and 17 Earth-masses. This is probably coincidence as there is no geophysical barrier to the
formation of intermediary bodies (see for instance OGLE-2005-BLG-390Lb) and we should expect to find planets
throughout the galaxy between two and twelve Earth-masses. If the star system is otherwise favourable, such
planets would be good candidates for life as they would be large enough to remain internally dynamic and
atmosphere retentive over billions of years but not so large as to accrete the gaseous shell which limits the
possibility of life formation.
• Note g: According to prevailing theory, the formation of the Moon commenced when a Mars-sized body struck
the Earth in a glancing collision late in its formation, and the ejected material coalesced and fell into orbit (see
giant impact hypothesis). In Rare Earth Ward and Brownlee emphasize that such impacts ought to be rare,
reducing the probability of other Earth-Moon type systems and hence the probability of other habitable planets.
Other moon formation processes are possible, however, and the proposition that a planet may be habitable in the
absence of a moon has not been disproven.
Further reading
• Abstracts from the Astrobiology Science Conference 2004 [56].
• Cohen, Jack and Ian Stewart. Evolving the Alien: The Science of Extraterrestrial Life, Ebury Press, 2002. ISBN
0-09-187927-2
• Dole, Stephen H. (1965). Habitable Planets for Man [57] (1st ed.). Rand Corporation. ISBN 0-444-00092-5.
• Fogg, Martyn J., ed. "Terraforming" (entire special issue) Journal of the British Interplanetary Society, April
1991
• Fogg, Martyn J. Terraforming: Engineering Planetary Environments, SAE International, 1995. ISBN
1-56091-609-5
• Gonzalez, Guillermo and Richards, Jay W. The Privileged Planet, Regnery, 2004. ISBN 0-89526-065-4
• Grinspoon, David. Lonely Planets: The Natural Philosophy of Alien Life, HarperCollins, 2004.
• Lovelock, James. Gaia: A New Look at Life on Earth. ISBN 0-19-286218-9
• Schmidt, Stanley and Robert Zubrin, eds. Islands in the Sky, Wiley, 1996. ISBN 0-471-13561-5
• Ward, Peter and Donald Brownlee. Rare Earth: Why Complex Life is Uncommon in the Universe, Springer, 2000.
ISBN 0-387-98701-0
• Webb, Stephen If The Universe Is Teeming With Aliens ... Where Is Everybody? Fifty Solutions to the Fermi
Paradox and the Problem of Extraterrestrial Life New York: January 2002 Springer-Verlag ISBN
978-0-387-95501-8
External links
• Alan Boss research papers [58]
• David Darling encyclopedia [59]
• General interest astrobiology [60]
• James Kasting research papers [61]
• Margaret Turnbull HabCat related files [62]
• Sol Station [63]
• Martyn J. Fogg Terraforming Information Pages [64]
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[42] Cain, Fraser; and Gay, Pamela (2007). "AstronomyCast episode 40: American Astronomical Society Meeting, May 2007" (http:/ /
media-c02m01. libsyn. com/ podcasts/ c50d001e8872db18d96cd44a73adccdc/ 46762eec/ astronomycast/ AstroCast-070611. mp3). Universe
Today. . Retrieved 2007-06-17.
[43] University of Washington (January 13, 2003). "'The end of the world' has already begun, UW scientists say" (http:/ / www. washington. edu/
newsroom/ news/ 2003archive/ 01-03archive/ k011303a. html). Press release. . Retrieved 2007-06-05.
[44] "M Dwarfs: The Search for Life is On, Interview with Todd Henry" (http:/ / www. astrobio. net/ news/ modules. php?op=modload&
name=News& file=article& sid=1694). Astrobiology Magazine. August 29, 2005. . Retrieved 2007-08-05..
[45] Mullen, Leslie (May 18, 2001). "Galactic Habitable Zones" (http:/ / www. astrobio. net/ news/ modules. php?op=modload& name=News&
file=article& sid=139). Astrobiology Magazine. . Retrieved 2007-08-05.
[46] Rare Earth, pp. 26–29.
[47] Dorminey, Bruce (July 2005). "Dark Threat". Astronomy: 40–45.
[48] Drake, Frank (1973). "Life on a Neutron Star". Astrobiology 1 (5): 5.
[49] Darling, David. "Neutron star, life on" (http:/ / www. daviddarling. info/ encyclopedia/ N/ neutronstarlife. html). The Encyclopedia of
Astrobiology, Astronomy, and Spaceflight. . Retrieved 2009-09-05.
[50] Sagan, C.; Salpeter, E. E. (1976). "Particles, environments, and possible ecologies in the Jovian atmosphere". The Astrophysical Journal
Supplement Series 32: 633–637. doi:10.1086/190414.
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Astronomy, and Spaceflight. . Retrieved 2007-08-06.
[52] Bortman, Henry (September 29, 2004). "Coming Soon: "Good" Jupiters" (http:/ / www. astrobio. net/ news/ modules. php?op=modload&
name=News& file=article& sid=1222). Astrobiology Magazine. . Retrieved 2007-08-05.
[53] Lunine, Jonathan I. (January 30, 2001). "The occurrence of Jovian planets and the habitability of planetary systems" (http:/ / www. pnas.
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Astrobiology Magazine. September 22, 2005. . Retrieved 2007-08-06.
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Abiogenesis
In natural science, abiogenesis
(pronounced /ˌeɪbaɪ.ɵˈdʒɛnɨsɪs/, AY-bye-oh-JEN-ə-siss) or
biopoesis is the study of how life on Earth could have
arisen from inanimate matter. It should not be confused
with evolution, which is the study of how groups of
already living things change over time, or with
cosmogony, which covers how the universe might have
arisen. Most amino acids, often called "the building
blocks of life", can form via natural chemical reactions
unrelated to life, as demonstrated in the Miller–Urey
experiment and similar experiments, which involved
Pre-Cambrian stromatolites in the Siyeh Formation, Glacier National
simulating some of the conditions of the early Earth, in
Park. In 2002, William Schopf of UCLA published a paper in the
a scientific laboratory.[1] In all living things, these scientific journal Nature arguing that geological formations such as
amino acids are organized into proteins, and the this possess 3.5 Ga (billion years old) fossilized cyanobacteria
construction of these proteins is mediated by nucleic microbes. If true, they would be evidence of the earliest known life
on earth.
acids. Which of these organic molecules first arose and
how they formed the first life is the focus of
abiogenesis.
In any theory of abiogenesis, two aspects of life have to be accounted for: replication, and metabolism. The question
of which came first gave rise to different types of theories. In the beginning, metabolism-first theories (Oparin
coacervate) were proposed, and only later thinking gave rise to modern, replication-first approach.
In modern, still somewhat limited understanding, the first living things on Earth are thought to be single cell
prokaryotes (which lack a cell nucleus), perhaps evolved from protobionts (organic molecules surrounded by a
Abiogenesis 19
membrane-like structure).[2] The oldest ancient fossil microbe-like objects are dated to be 3.5 Ga (billion years old),
approximately one billion years after the formation of the Earth itself.[3] [4] By 2.4 Ga, the ratio of stable isotopes of
carbon, iron and sulfur shows the action of living things on inorganic minerals and sediments[5] [6] and molecular
biomarkers indicate photosynthesis, demonstrating that life on Earth was widespread by this time.[7] [8]
The sequence of chemical events that led to the first nucleic acids is not known. Several hypotheses about early life
have been proposed, most notably the iron-sulfur world theory (metabolism without genetics) and the RNA world
hypothesis (RNA life-forms).
Conceptual history
Spontaneous generation
Until the early 19th century, people generally believed in the ongoing spontaneous generation of certain forms of life
from non-living matter. This was paired with heterogenesis, the belief that one form of life derives from a different
form (e.g. bees from flowers).[9] Classical notions of abiogenesis, now more precisely known as spontaneous
generation, held that certain complex, living organisms are generated by decaying organic substances. According to
Aristotle it was a readily observable truth that aphids arise from the dew which falls on plants, fleas from putrid
matter, mice from dirty hay, crocodiles from rotting logs at the bottom of bodies of water, and so on.[10]
In the 17th century, such assumptions started to be questioned; for example, in 1646, Sir 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 conclusions were not widely accepted. For example, his
contemporary, Alexander Ross wrote: "To question this (i.e., spontaneous generation) is to question reason, sense
and experience. If he doubts of this let him go to Egypt, and there he will find the fields swarming with mice, begot
of the mud of Nylus, to the great calamity of the inhabitants."[11]
In 1665, Robert Hooke published the first drawings of a microorganism. Hooke was followed in 1676 by Anton van
Leeuwenhoek, who drew and described microorganisms that are now thought to have been protozoa and bacteria.[12]
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.
The first solid evidence against spontaneous generation came in 1668 from Francesco Redi, who proved 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 seemed to be biogenesis: that every living thing came from a pre-existing living thing (omne
vivum ex ovo, Latin for "every living thing from an egg").
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 which demonstrated that organisms such as bacteria and fungi
do not spontaneously appear in sterile, nutrient-rich media.
Abiogenesis 20
Pasteur and Darwin

By the middle of the 19th century, the theory of biogenesis had accumulated so much
evidential support, due to the work of Louis Pasteur and others, that the alternative
theory of spontaneous generation had been effectively disproven. Pasteur himself
remarked, after a definitive finding in 1864, "Never will the doctrine of spontaneous
generation recover from the mortal blow struck by this simple experiment."[13] The
collapse of spontaneous generation, however, left a vacuum of scientific thought on
the question of how life had first arisen.
In a letter to Joseph Dalton Hooker on February 1, 1871,[14] Charles Darwin

addressed the question, suggesting that the original spark of life may have begun in a
"warm little pond, with all sorts of ammonia and phosphoric salts, lights, heat, Charles Darwin in 1879.
electricity, etc. present, so that a protein 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."[15] In other words,
the presence of life itself makes the search for the origin of life dependent on the sterile conditions of the laboratory.
"Primordial soup" theory

No new notable research or theory on the subject appeared until 1924,
when Alexander Oparin reasoned that atmospheric oxygen prevents the
synthesis of certain organic compounds that are necessary building blocks
for the evolution of life. In his The Origin of Life,[16] [17] Oparin proposed
that the "spontaneous generation of life" that had been attacked by Louis
Pasteur, did in fact occur once, but was now impossible because the
conditions found in the early earth had changed, and the presence of living
organisms would immediately consume any spontaneously generated
organism. Oparin argued that a "primeval soup" of organic molecules could
be created in an oxygen-less atmosphere through the action of sunlight.
These would combine in ever-more complex fashions 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 those factors which promote "cell
integrity" survive, those that do not become extinct. Many modern theories
of the origin of life still take Oparin's ideas as a starting point.
Alexander Oparin (right) at the laboratory.
Around the same time, J. B. S. Haldane suggested that the Earth's pre-biotic
oceans–very different from their modern counterparts–would have formed a "hot dilute soup" in which organic
compounds could have formed. This idea was called biopoiesis or biopoesis, the process of living matter evolving
from self-replicating but nonliving molecules.[18] [19]
Early conditions
Morse and MacKenzie have suggested that oceans may have appeared first in the Hadean eon, as soon as two
hundred million years (200 Ma) after the Earth was formed, in a hot 100 °C (212 °F) reducing environment, and that
the pH of about 5.8 rose rapidly towards neutral.[20] This has been supported by Wilde[3] who has pushed the date of
the zircon crystals found in the metamorphosed quartzite of Mount Narryer in Western Australia, previously thought
to be 4.1–4.2 Ga, to 4.404 Ga. This means that oceans and continental crust existed within 150 Ma of Earth's
Abiogenesis 21
formation.
Despite this, the Hadean environment was one highly hazardous to life. Frequent collisions with large objects, up to
500 kilometres (310 mi) in diameter, would have been sufficient to vaporise the ocean within a few months of
impact, with hot steam mixed with rock vapour leading to high altitude clouds completely covering the planet. After
a few months the height of these clouds would have begun to decrease but the cloud base would still have been
elevated for about the next thousand years. After that, it would have begun to rain at low altitude. For another two
thousand years rains would slowly have drawn down the height of the clouds, returning the oceans to their original
depth only 3,000 years after the impact event.[21]
Between 3.8 and 4.1 Ga, changes in the orbits of the gaseous giant planets may have caused a late heavy
bombardment that pockmarked the moon and other inner planets (Mercury, Mars, and presumably Earth and Venus).
This would likely have sterilized the planet had life appeared before that time.
By examining the time interval between such devastating environmental events, the time interval when life might
first have come into existence can be found for different early environments. The study by Maher and Stevenson
shows that if the deep marine hydrothermal setting provides a suitable site for the origin of life, abiogenesis could
have happened as early as 4.0 to 4.2 Ga, whereas if it occurred at the surface of the earth abiogenesis could only have
occurred between 3.7 and 4.0 Ga.[22]
Other research suggests a colder start to life. Work by Leslie Orgel and colleagues on the synthesis of purines has
shown that freezing temperatures are advantageous, due to the concentrating effect for key precursors such as
HCN.[23] Research by Stanley Miller and colleagues suggested that while adenine and guanine require freezing
conditions for synthesis, cytosine and uracil may require boiling temperatures.[24] Based on this research, Miller
suggested a beginning of life involving freezing conditions and exploding meteorites.[25] An article in Discover
Magazine points to research by the Miller group indicating the formation of seven different amino acids and 11 types
of nucleobases in ice when ammonia and cyanide were left in a freezer from 1972–1997.[26] [27] This article also
describes research by Christof Biebricher showing the formation of RNA molecules 400 bases long under freezing
conditions using an RNA template, a single-strand chain of RNA that guides the formation of a new strand of RNA.
As that new RNA strand grows, it adheres to the template.[28] 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.
Evidence of the early appearance of life comes from the Isua supercrustal belt in Western Greenland and from
similar formations in the nearby Akilia Islands. Carbon entering into rock formations has a ratio of Carbon-13 (13C)
to Carbon-12 (12C) of about −5.5 (in units of δ13C), where because of a preferential biotic uptake of 12C, biomass
has a δ13C of between −20 and −30. These isotopic fingerprints are preserved in the sediments, and Mojzis has used
this technique to suggest that life existed on the planet already by 3.85 billion years ago.[29] Lazcano and Miller
(1994) suggest that the rapidity of the evolution of life is dictated by the rate of recirculating water through
mid-ocean submarine vents. Complete recirculation takes 10 million years, thus any organic compounds produced by
then would be altered or destroyed by temperatures exceeding 300 °C (572 °F). They estimate that the development
of a 100 kilobase genome of a DNA/protein primitive heterotroph into a 7000 gene filamentous cyanobacterium
would have required only 7 Ma.[30]
Abiogenesis 22
Current models
There is no truly "standard model" of the origin of life. Most currently accepted models draw at least some elements
from the framework laid out by the Oparin-Haldane hypothesis. Under that umbrella, however, are a wide array of
disparate discoveries and conjectures such as the following, listed in a rough order of postulated emergence:
1. Some theorists suggest that the atmosphere of the early Earth may have been chemically reducing in nature,
composed primarily of methane (CH4), ammonia (NH3), water (H2O), hydrogen sulfide (H2S), carbon dioxide
(CO2) or carbon monoxide (CO), and phosphate (PO43-), with molecular oxygen (O2) and ozone (O3) either rare
or absent.
2. In such a reducing atmosphere, electrical activity can catalyze the creation of certain basic small molecules
(monomers) of life, such as amino acids. This was demonstrated in the Miller–Urey experiment by Stanley L.
Miller and Harold C. Urey in 1953.
3. Phospholipids (of an appropriate length) can spontaneously form lipid bilayers, a basic component of the cell
membrane.
4. A fundamental question is about the nature of the first self-replicating molecule. Since replication is
accomplished in modern cells through the cooperative action of proteins and nucleic acids, the major schools of
thought about how the process originated can be broadly classified as "proteins first" and "nucleic acids first".
5. The principal thrust of the "nucleic acids first" argument is as follows:
1. The polymerization of nucleotides into random RNA molecules might have resulted in self-replicating
ribozymes (RNA world hypothesis)
2. Selection pressures for catalytic efficiency and diversity might have resulted in ribozymes which catalyse
peptidyl transfer (hence formation of small proteins), since oligopeptides complex with RNA to form better
catalysts. The first ribosome might have been created by such a process, resulting in more prevalent protein
synthesis.
3. Synthesized proteins might then outcompete ribozymes in catalytic ability, and therefore become the dominant
biopolymer, relegating nucleic acids to their modern use, predominantly as a carrier of genomic information.
As of 2010, no one has yet synthesized a "protocell" using basic components which would have the necessary
properties of life (the so-called "bottom-up-approach"). Without such a proof-of-principle, explanations have tended
to be short on specifics. However, some researchers are working in this field, notably Steen Rasmussen at Los
Alamos National Laboratory and Jack Szostak at Harvard University. Others have argued that a "top-down
approach" is more feasible. One such approach, attempted by Craig Venter and others at The Institute for Genomic
Research, involves engineering existing prokaryotic cells with progressively fewer genes, attempting to discern at
which point the most minimal requirements for life were reached. The biologist John Desmond Bernal coined the
term Biopoesis for this process , and suggested that there were a number of clearly defined "stages" that could be
recognised in explaining the origin of life.
• Stage 1: The origin of biological monomers
• Stage 2: The origin of biological polymers
• Stage 3: The evolution from molecules to cell
Bernal suggested that evolution may have commenced early, some time between Stage 1 and 2[31] .
Abiogenesis 23
Origin of organic molecules

There are two possible sources of organic molecules on the early Earth:
1. Terrestrial origins–organic synthesis driven by impact shocks or by other energy sources (such as ultraviolet light
or electrical discharges) (eg.Miller's experiments)
2. Extraterrestrial origins–delivery by objects (e.g. carbonaceous chondrites) or gravitational attraction of organic
molecules or primitive life-forms from space
Recently, estimates of these sources suggest that the heavy bombardment before 3.5 Ga within the early atmosphere
made available quantities of organics comparable to those produced by other energy sources.[32]
"Soup" theory today: Miller's experiment and subsequent work

Biochemist Robert Shapiro has summarized the "Primordial Soup" theory of Oparin and Haldane in its "mature
form" as follows:[33]
1. The early Earth had a chemically reducing atmosphere, as discussed above.
2. This atmosphere, exposed to energy in various forms, produced simple organic compounds ("monomers").
3. These compounds accumulated in a "soup", which may have been concentrated at various locations (Shorelines,
oceanic vents etc.).
4. By further transformation, more complex organic polymers— and ultimately life— developed in the soup.
Regarding the reducing atmosphere

Whether the mixture of gases used in the Miller–Urey experiment truly reflects the atmospheric content of early
Earth is a controversial topic. Other less reducing gases produce a lower yield and variety. It was once thought that
appreciable amounts of molecular oxygen were present in the prebiotic atmosphere , which would have essentially
prevented the formation of organic molecules; however, the current scientific consensus is that such was not the
case. (See Oxygen catastrophe).
Regarding monomer formation

One of the most important pieces of experimental support for the "soup" theory came in 1953. A graduate student,
Stanley Miller, and his professor, 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 reduced mixture of
gases–methane, ammonia and hydrogen–to form basic organic monomers, such as amino acids.[34] This provided
direct experimental support for the second point of the "soup" theory as described above, and it is around the
remaining two points of the theory that much of the debate now centers.
Apart from the Miller–Urey experiment, described above, the next most important step in research on prebiotic
organic synthesis was the demonstration by John Oró that the nucleic acid purine base, adenine, was formed by
heating aqueous ammonium cyanide solutions.[35] In support of abiogenesis in eutectic ice (see above), more recent
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).[36]
Abiogenesis 24
Regarding monomer accumulation

The "soup" theory relies on the assumption proposed by Darwin (see above) that in an environment with no
pre-existing life, organic molecules may have accumulated and provided an environment for chemical evolution.
Regarding further transformation

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 formed in high concentration during the
Miller–Urey and Oró experiments. The Miller experiment, for example, produces many substances that would
undergo cross-reactions with the amino acids or terminate the peptide chain.
More fundamentally, it can be argued that the most crucial challenge unanswered by this theory is how the relatively
simple organic building blocks polymerise and form more complex structures, interacting in consistent ways to form
a protocell. For example, in an aqueous environment hydrolysis of oligomers/polymers into their constituent
monomers would be favored over the condensation of individual monomers into polymers.
The deep sea vent theory

The deep sea vent, or hydrothermal vent, theory for the origin of life on Earth posits that life may have begun at
submarine hydrothermal vents, where hydrogen-rich fluids emerge from below the sea floor and interface with
carbon dioxide-rich ocean water. Sustained chemical energy in such systems is derived from redox reactions, in
which electron donors, such as molecular hydrogen, react with electron acceptors, such as carbon dioxide (see
iron-sulfur world theory).
Fox's experiments
In the 1950s and 1960s, Sidney W. Fox studied the spontaneous formation of peptide structures under conditions that
might plausibly have existed early in Earth's history. He demonstrated that amino acids could spontaneously form
small peptides. These amino acids and small peptides could be encouraged to form closed spherical membranes,
called protenoid microspheres, which show many of the basic characteristics of 'life'.[37]
Eigen's hypothesis
In the early 1970s the problem of the origin of life was approached by Manfred Eigen and Peter Schuster of the Max
Planck Institute for Biophysical Chemistry. They examined the transient stages between the molecular chaos and a
self-replicating hypercycle in a prebiotic soup.[38]
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 that RNA, in certain circumstances,
forms itself into ribozymes, capable of catalyzing their own chemical reactions.[39] However, these reactions are
limited to self-excisions (in which a longer RNA molecule becomes shorter), and much rarer small additions that are
incapable of coding for any useful protein. The hypercycle theory is further degraded since the hypothetical RNA
would require the existence of complex biochemicals such as nucleotides which are not formed under the conditions
proposed by the Miller–Urey experiment.
Abiogenesis 25
Hoffmann's contributions
Geoffrey W. Hoffmann, a student of Eigen, contributed to the concept of life involving both replication and
metabolism emerging from catalytic noise. His contributions included showing that an early sloppy translation
machinery can be stable against an error catastrophe of the type that had been envisaged as problematical by Leslie
Orgel ("Orgel's paradox")[40] [41] and calculations regarding the occurrence of a set of required catalytic activities
together with the exclusion of catalytic activities that would be disruptive. This is called the stochastic theory of the
origin of life.[42]
Wächtershäuser's hypothesis
Another possible answer to this polymerization conundrum was

provided in 1980s by the German chemist Günter Wächtershäuser, in
his iron-sulfur world theory. In this theory, he postulated the evolution
of (bio)chemical pathways as fundamentals of the evolution of life.
Moreover, he presented a consistent system of tracing today's
biochemistry back to ancestral reactions that 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 (such as simulated lightning or UV
irradiation), "Wächtershäuser systems" come with a built-in source of
energy, sulfides of iron and other minerals (e.g. pyrite). The energy
released from redox reactions of these metal sulfides is not only
available for the synthesis of organic molecules, but also for the
formation of oligomers and polymers. It is therefore hypothesized that
such systems may be able to evolve into autocatalytic sets of
self-replicating, metabolically active entities that would predate the life Deep-sea black smoker
forms known today.
The experiment produced a relatively small yield of dipeptides (0.4% to 12.4%) and a smaller yield of tripeptides
(0.10%) but the authors also noted that: "under these same conditions dipeptides hydrolysed rapidly."[43]
Radioactive beach hypothesis

Zachary Adam at the University of Washington, Seattle, claims that stronger tidal processes from a much closer
moon 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.[44] According to computer
models reported in Astrobiology,[45] 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 provides
sufficient energy to generate organic molecules, such as amino acids and sugars from acetonitrile in water.
Radioactive monazite also releases soluble phosphate into regions between sand-grains, making it biologically
"accessible". Thus amino acids, sugars and soluble phosphates can all be simultaneously produced, according to
Adam. Radioactive actinides, then in greater concentrations, could have formed part of organo-metallic complexes.
These complexes could have been important early catalysts to living processes.
John Parnell of the University of Aberdeen suggests that such a process could provide part of the "crucible of life" on
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 believed to have had many smaller "platelets" it
would provide a suitable environment for such processes.[46]
Abiogenesis 26
Thermodynamic Origin of Life: Ultraviolet and Temperature-Assisted Replication (UVTAR) Model

Karo Michaelian of the National Autonomous University of Mexico (UNAM) points out that any model for the
origin of life must take into account the fact that life is an irreversible thermodynamic process which arises and
persists to produce entropy. Entropy production is not incidental to the process of life, but rather the fundamental
reason for its existence. Present day life augments the entropy production of Earth by catalysing the water cycle
through evapotranspiration.[47] Michaelian argues that if the thermodynamic function of life today is to produce
entropy through coupling with the water cycle, 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 200 nm - 300 nm wavelength range, just that high energy part of the sun's spectrum that
could have penetrated the dense prebiotic atmosphere. Cnossen et al.[48] have shown that the amount of UV light
reaching the Earth's surface in the Archean could have been up to 31 orders of magnitude larger than it is today at
260 nm where RNA and DNA absorb most strongly. Absorption and dissipation of UV light by these organic
molecules at the Archean ocean surface would have increased significantly the temperature of the surface skin layer
leading to enhanced evaporation and thus augmenting the primitive water cycle. Since absorption and dissipation of
high energy photons is an entropy producing process, Michaelian argues that non-equilbrium abiogenic synthesis of
RNA and DNA utilizing UV light [49] would have been thermodynamically favored.
A simple mechanism to explain the replication of RNA and DNA without the use of enzymes can also be given
within the same thermodynamic framework by assuming that life arose when the temperature of the primitive seas
had cooled to somewhat below the denaturing temperature of RNA or DNA (based on the ratio of 18O/16O found in
cherts of the Barberton greenstone belt of South Africa of about 3.5 to 3.2 Ga., surface temperatures are predicted to
have been around 70±15 °C [50] , similar to RNA or DNA denaturing temperatures). During the night, the surface
water temperature would be below the denaturing temperature and single strand RNA/DNA could act as a template
for the formation of double strand RNA/DNA. During the daylight hours, RNA and DNA would absorb UV light
and convert this directly to heating of the ocean surface, raising the local temperature enough to allow for denaturing
of RNA and DNA. The copying process would be repeated during the cool period overnight [51] . Such a temperature
assisted mechanism of replication bears similarity to Polymerase Chain Reaction (PCR), a routine laboratory
procedure to multiply DNA segments. Michaelian suggests that traditional origin of life research, expecting to
describe the emergence of life from near-equilibrium conditions, is erroneous and that non-equilibrium conditions
must be considered, in particular, the importance of entropy production to the emergence of life.
Since denaturation would be most probable in the late afternoon when the Archean sea surface temperature would be
highest, and since late afternoon sunlight is somewhat circularly polarized, the homochirality of the organic
molecules of life can also be explained within the proposed thermodynamic framework.[52]
Models to explain homochirality

Some process in chemical evolution must account for the origin of homochirality, i.e. all building blocks in living
organisms having the same "handedness" (amino acids being left-handed, nucleic acid sugars (ribose and
deoxyribose) being right-handed, and chiral phosphoglycerides). 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. This is called
a racemic mixture. Clark has suggested that homochirality may have started in space, as the studies of the amino
acids on the Murchison meteorite showed L-alanine to be more than twice as frequent as its D form, and L-glutamic
acid was more than 3 times prevalent than its D counterpart. It is suggested that polarised light has the power to
destroy one enantiomer within the proto-planetary disk. Noyes[53] showed that beta decay caused the breakdown of
D-leucine, in a racemic mixture, and that the presence of 14C, present in larger amounts in organic chemicals in the
early Earth environment, could have been the cause. Robert M. Hazen reports upon experiments conducted in which
various chiral crystal surfaces act as sites for possible concentration and assembly of chiral monomer units into
macromolecules.[54] Once established, chirality would be selected for.[55] Work with organic compounds found on
meteorites tends to suggest that chirality is a characteristic of abiogenic synthesis, as amino acids show a left-handed
Abiogenesis 27
bias, whereas sugars show a predominantly right-handed bias.[56]
Self-organization and replication

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. For example Martin and
Russel show that physical compartmentation by cell membranes from the environment and self-organization of
self-contained redox reactions are the most conserved attributes of living things, and they argue therefore that
inorganic matter with such attributes would be life's most likely last common ancestor.[57]
Virus self-assembly within host cells has implications for the study of the origin of life,[58] as it lends further
credence to the hypothesis that life could have started as self-assembling organic molecules.[59] [60]
From organic molecules to protocells

The question "How do simple organic molecules form a protocell?" is largely unanswered but there are many
hypotheses. Some of these postulate the early appearance of nucleic acids ("genes-first") whereas others postulate the
evolution of biochemical reactions and pathways first ("metabolism-first"). Recently, trends are emerging to create
hybrid models that combine aspects of both.
"Genes first" models: the RNA world

The RNA world hypothesis describes an early Earth with self-replicating and catalytic RNA but no DNA or proteins.
This has spurred scientists to try to determine if relatively short RNA molecules could have spontaneously formed
that were capable of catalyzing their own continuing replication.[61] A number of hypotheses of modes of formation
have been put forward.[62] Early cell membranes could have formed spontaneously from proteinoids, protein-like
molecules that are produced when amino acid solutions are heated–when present at the correct concentration in
aqueous solution, these form microspheres which are observed to behave similarly to membrane-enclosed
compartments. Other possibilities include systems of chemical reactions taking place within clay substrates or on the
surface of pyrite rocks. Factors supportive of an important role for RNA in early life include its ability to act both to
store information and catalyse chemical reactions (as a ribozyme); its many important roles as an intermediate in the
expression 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 molecule under conditions approximating the early Earth.
Relatively short RNA molecules which can duplicate others have been artificially produced in the lab.[63] Such
replicase RNA, which functions as both code and catalyst provides a template upon which copying can occur. Jack
Szostak has shown that certain catalytic RNAs can, indeed, join smaller RNA sequences together, creating the
potential, in the right conditions for self-replication. If these were present, Darwinian selection would favour the
proliferation of such self-catalysing structures, to which further functionalities could be added.[64] Lincoln and Joyce
identified an RNA enzyme capable of self sustained replication.[65]
Researchers have pointed out difficulties for the abiotic synthesis of nucleotides from cytosine and uracil.[66]
Cytosine has a half-life of 19 days at 100 °C (212 °F) and 17,000 years in freezing water.[67] Larralde et al., say that
"the generally accepted prebiotic synthesis of ribose, the formose reaction, yields numerous sugars without any
selectivity."[68] 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.[69]
A slightly different version of the RNA-world hypothesis is that a different type of nucleic acid, such as PNA, TNA
or GNA, was the first one to emerge as a self-reproducing molecule, to be replaced by RNA only later.[70] [71]
Pyrimidine ribonucleosides and their respective nucleotides have been prebiotically synthesised by a sequence of
reactions which by-pass the free sugars, and are assembled in a stepwise fashion by going against the dogma that
nitrogenous and oxygenous chemistries should be avoided. In a series of publications, The Sutherland Group at the
Abiogenesis 28
School of Chemistry, University of Manchester have 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.[72] This can
be viewed as a prebiotic purification step, where the said compound spontaneously crystallised 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.[73] 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 polymerise into RNA. This paper also highlights the possibility for the
photo-sanitization of the pyrimidine-2',3'-cyclic phosphates.[49] James Ferris's studies have shown that clay minerals
of montmorillonite will catalyze the formation of RNA in aqueous solution, by joining activated mono RNA
nucleotides to join together to form longer chains.[74] Although these chains have random sequences, the possibility
that one sequence began to non-randomly increase its frequency by increasing the speed of its catalysis is possible to
"kick start" biochemical evolution.
"Metabolism first" models

Several models reject the idea of the self-replication of a "naked-gene" and postulate the emergence of a primitive
metabolism which could provide an environment for the later emergence of RNA replication.
Iron-sulfur world
One of the earliest incarnations of this idea was put forward in 1924 with Alexander Oparin's notion of primitive
self-replicating vesicles which predated the discovery of the structure of DNA. More recent variants in the 1980s and
1990s include Günter 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 in that decade.
However, the idea that a closed metabolic cycle, such as the reductive citric acid cycle, could form spontaneously
(proposed by Günter Wächtershäuser) remains debated. In an article entitled "Self-Organizing Biochemical
Cycles",[75] the late Leslie Orgel summarized his analysis of the proposal 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." 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
recognised ways of carbon dioxide fixation in nature today) would be compatible with the idea of self-organisation
on a metal sulfide surface. The key enzyme of this pathway, carbon monoxide dehydrogenase/acetyl-CoA synthase
harbours mixed nickel-iron-sulfur clusters in its reaction centers and catalyses the formation of acetyl-CoA (which
may be regarded as a modern form of acetyl-thiol) in a single step.
Thermosynthesis world
Today’s bioenergetic process of fermentation is related to the just mentioned citric acid cycle or the Acetyl-CoA
pathway that have been connected to the primordial iron-sulfur world. In a different approach, today’s bioenergetic
process of chemiosmosis, which plays an essential role in cellular respiration and photosynthesis, is considered as
more fundamental than fermentation: in Anthonie Muller’s “thermosynthesis world” the ATP Synthase enzyme that
sustains chemiosmosis is proposed as today’s enzyme that is the closest connected to the first metabolic process.[76]
[77]
First life needed an energy source to bring about the 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
Abiogenesis 29
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 after a temperature change was
released upon a thermal unfolding.
The energy source of the thermosynthesis world was thermal cycling, the result of suspension of the protocell 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. In the beginning this First Protein also synthesized a library with many proteins, of which only
a minute fraction had thermosynthesis capabilities. Just as proposed by Dyson [78] for the first proteins, the First
Protein propagated functionally: it made daughters with similar capabilities, but it did not copy itself. Functioning
daughters consisted of different amino acid sequences.
Over a long time, RNA sequences were selected among the at first randomly synthesized RNAs by the criterion of
speed and efficiency increase of First Protein synthesis, for instance by the creation of RNA that functioned as
messenger RNA[79] , Transfer RNA[80] and ribosomal RNA, or, even more generally, all the components of the RNA
World were also generated and selected. The thermosynthesis world therefore in theory accounts for the origin of the
genetic machinery.
Whereas the iron-sulfur world identifies a circular pathway as the most simple—and therefore assumes the existence
of enzymes—the thermosynthesis world does not even invoke a pathway, and does not assume the existence of
regular enzymes: ATP Synthase’s binding change mechanism resembles a physical adsorption process that yields
free energy[81] , rather than a regular enzyme’s mechanism, which decreases the free energy. The RNA World also
implies the existence of several enzymes. But even the emergence of a single enzyme by chance is implausible.[82]
The thermosynthesis world is therefore more simple, and thus more plausible, than the iron-sulfur and RNA worlds.
Possible role of bubbles

Waves breaking on the shore create a delicate foam composed of bubbles. Winds sweeping across the ocean have a
tendency to drive things to shore, much like driftwood collecting on the beach. It is possible that organic molecules
were concentrated on the shorelines in much the same way. Shallow coastal waters also tend to be warmer, further
concentrating the molecules through evaporation. While bubbles composed mostly of water burst quickly, water
containing amphiphiles forms much more stable bubbles, lending more time to the particular bubble to perform these
crucial reactions.
Amphiphiles are oily compounds containing a hydrophilic head on one or both ends of a hydrophobic molecule.
Some amphiphiles have the tendency to spontaneously form membranes in water. A spherically closed membrane
contains water and is a hypothetical precursor to the modern cell membrane. If a protein would increase the integrity
of its parent bubble, that bubble had an advantage, and was placed at the top of the natural selection waiting list.
Primitive reproduction can be envisioned when the bubbles burst, releasing the results of the 'experiment' into the
surrounding medium. Once enough of the 'right stuff' was released into the medium, the development of the first
prokaryotes, eukaryotes, and multicellular organisms could be achieved.[83]
Similarly, bubbles formed entirely out of protein-like molecules, called microspheres, will form spontaneously under
the right conditions. But they are not a likely precursor to the modern cell membrane, as cell membranes are
composed primarily of lipid compounds rather than amino-acid compounds (for types of membrane spheres
associated with abiogenesis, see protobionts, micelle, coacervate).
A recent model by Fernando and Rowe[84] suggests that the enclosure of an autocatalytic non-enzymatic metabolism
within protocells may have been one way of avoiding the side-reaction problem that is typical of metabolism first
models.
Abiogenesis 30
Other models
Autocatalysis
In 1993 Stuart Kauffman proposed that life initially arose as autocatalytic chemical networks.[85]
British ethologist Richard Dawkins wrote about autocatalysis as a potential explanation for the origin of life in his
2004 book The Ancestor's Tale. Autocatalysts are substances which catalyze the production of themselves, and
therefore have the property of being a simple molecular replicator. In his book, Dawkins cites experiments
performed by Julius Rebek and his colleagues at the Scripps Research Institute in California in which they combined
amino adenosine and pentafluorophenyl ester with the autocatalyst amino adenosine triacid ester (AATE). One
system from the experiment contained variants of AATE which catalysed 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.
Clay theory
A model for the origin of life based on clay was forwarded by A. Graham Cairns-Smith of the University of Glasgow
in 1985 and explored as a plausible illustration by several other scientists, including Richard Dawkins[86] . Clay
theory postulates that complex organic molecules arose gradually on a pre-existing, non-organic replication
platform—silicate crystals in solution. Complexity in companion molecules developed as a function of selection
pressures on types of clay crystal is then exapted to serve the replication of organic molecules independently of their
silicate "launch stage".
Cairns-Smith is a staunch critic of other models of chemical evolution.[87] However, he admits that like many models
of the origin of life, his own also has its shortcomings (Horgan 1991).
In 2007, Kahr and colleagues reported their experiments to examine 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 crystal system and found that the imperfections in the mother crystals were indeed reproduced in
the daughters. The daughter crystals had many additional imperfections. For a gene-like behavior the additional
imperfections should be much less than the parent ones, thus Kahr concludes that the crystals "were not faithful
enough to store and transfer information from one generation to the next".[88] [89]
Gold's "Deep-hot biosphere" model

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. The discovery in the late 1990s of nanobes (filamental structures that are smaller than
bacteria, but that may contain DNA) in deep rocks [90] might be seen as lending support to Gold's theory.
It is now reasonably well established that microbial life is plentiful at shallow depths in the Earth, up to 5 kilometres
(3.1 mi) below the surface,[90] in the form of extremophile archaea, rather than the better-known eubacteria (which
live in more accessible conditions). It is claimed that discovery of microbial life below the surface of another body in
our solar system would lend significant credence to this theory. Thomas 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 flow of 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.
Abiogenesis 31
"Primitive" extraterrestrial life

An alternative to Earthly abiogenesis is the hypothesis that primitive life may have originally formed
extraterrestrially, either in space or on a nearby planet (Mars). (Note that exogenesis is related to, but not the same
as, the notion of panspermia). A supporter of this theory was Francis Crick.
Organic compounds are relatively common in space, especially in the outer solar system where volatiles are not
evaporated by solar heating.[91] Comets are encrusted by 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 irradiation by ultraviolet light. It is supposed that a rain of material from comets could have brought
significant quantities of such complex organic molecules to Earth.
An alternative but related hypothesis, proposed to explain the presence of life on Earth so soon after the planet had
cooled down, with apparently very little time for prebiotic evolution, is that life formed first on early Mars. Due to its
smaller size Mars cooled before Earth (a difference of hundreds of millions of years), allowing prebiotic processes
there while Earth was still too hot. Life was then transported to the cooled Earth when crustal material was blasted
off Mars by asteroid and comet impacts. Mars continued to cool faster and eventually became hostile to the
continued evolution or even existence of life (it lost its atmosphere due to low volcanism); Earth is following the
same fate as Mars, but at a slower rate.
Neither hypothesis actually answers the question of how life first originated, but merely shifts it to another planet or
a comet. However, the advantage of an extraterrestrial origin of primitive life is that life is not required to have
evolved on each planet it occurs on, but rather in a single location, and then spread about the galaxy to other star
systems via cometary and/or meteorite impact. Evidence to support the plausibility of the concept is scant, but it
finds support in recent study of Martian meteorites found in Antarctica and in studies of extremophile microbes.[92]
Additional support comes from a recent discovery of a bacterial ecosytem whose energy source is radioactivity.[93]
A 2001 experiment led by Jason Dworkin[94] subjected a frozen mixture of water, methanol, ammonia and carbon
monoxide to UV radiation, mimicking conditions found in an extraterrestrial environment. This combination yielded
large amounts of organic material that self-organised to form bubbles or micelles when immersed in water. Dworkin
considered these bubbles to resemble cell membranes that enclose and concentrate the chemistry of life, separating
their interior from the outside world.
The bubbles produced in these experiments were between 10 to 40 micrometres (0.00039 to 0.0016 in), or about the
size of red blood cells. Remarkably, the bubbles fluoresced, or glowed, when exposed to UV light. Absorbing UV
and converting it into visible light in this way was considered one possible way of providing energy to a primitive
cell. If such bubbles played a role in the origin of life, the fluorescence could have been a precursor to primitive
photosynthesis. Such fluorescence also provides the benefit of acting as a sunscreen, diffusing any damage that
otherwise would be inflicted by UV radiation. Such a protective function would have been vital for life on the early
Earth, since the ozone layer, which blocks out the sun's most destructive UV rays, did not form until after
photosynthetic life began to produce oxygen.[56]
Extraterrestrial amino acids

Another idea is that amino acids which were formed extra-terrestrially arrived on Earth via comets. In 2009 it was
announced by NASA that scientists have identified one of the fundamental chemical buildings blocks of life in a
comet for the first time: glycine, an amino acid, was detected in the material ejected from Comet Wild-2 in 2004 and
grabbed by NASA's Stardust probe. Tiny grains, just a few thousandths of a millimetre in size, were collected from
the comet and returned to Earth in 2006 in a sealed capsule, and distributed among the world's leading astro-biology
labs. NASA said in a statement that it took some time for the investigating team, led by Dr Jamie Elsila, to convince
itself that the glycine signature found in Stardust's sample bay was genuine and not just Earthly contamination.
Glycine has been detected in meteorites before and there are also observations in interstellar gas clouds claimed for
telescopes, but the Stardust find is described as a first in cometary material. It is known that prior to the emergence
Abiogenesis 32
of life on Earth, the early solar system's planets were regularly bombarded by comets. Dr. Carl Pilcher, who leads
NASA's 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."[95]
Lipid World
This theory postulates that the first self-replicating object was lipid-like.[96] It is known that phospholipids form
bilayers in water while under agitation– the same structure as in cell membranes. These molecules were not present
on early Earth, however 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 favorably. Still, no biochemical
mechanism has been offered to support the Lipid World theory.
Polyphosphates
The problem with 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.[97] [98] Polyphosphates are formed
by polymerization of ordinary monophosphate ions PO4−3. Several mechanisms for such polymerization have been
suggested. Polyphosphates cause polymerization of amino acids into peptides . They are also logical precursors in
the synthesis of such key biochemical compounds as 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.[99]
PAH world hypothesis

Other sources of complex molecules have been postulated, including extraterrestrial stellar or interstellar origin. For
example, from spectral analyses, organic molecules are known to be present in comets and meteorites. In 2004, a
team detected traces of polycyclic aromatic hydrocarbons (PAH's) in a nebula.[100] Those are the most complex
molecules so far found in space. The use of PAH's has also been proposed as a precursor to the RNA world in the
PAH world hypothesis.[101] The Spitzer Space Telescope has recently detected a star, HH 46-IR, which is forming
by a process similar to that by which the sun formed. In the disk of material surrounding the star, there is a very large
range of molecules, including cyanide compounds, hydrocarbons, and carbon monoxide. PAHs have also been found
all over the surface of galaxy M81, which is 12 million light years away from the Earth, confirming their widespread
distribution in space.[102]
Abiogenesis 33
Multiple genesis
Different forms of life may have appeared quasi-simultaneously in the early history of Earth.[103] The other forms
may be extinct, leaving distinctive fossils through their different biochemistry (e.g., using arsenic instead of
phosphorus), survive as extremophiles, or simply be unnoticed through their being analogous to organisms of the
current life tree. Hartman[104] for example combines a number of theories together, by proposing 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 biopoesis, 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.
Lynn Margulis's endosymbiotic theory suggests that multiple forms of bacteria entered into symbiotic relationship to
form the eukaryotic cell. The horizontal transfer of genetic material between bacteria promotes such symbiotic
relationships, and thus many separate organisms may have contributed to building what has been recognised as the
Last Universal Common Ancestor (LUCA) of modern organisms. James Lovelock's Gaia theory, proposes that such
bacterial symbiosis establishes the environment as a system produced by and supportive of life. His arguments
strongly weaken the case for life having evolved elsewhere in the solar system.
See also
• Astrochemistry
• Autocatalytic reactions and order creation
• Biogenesis
• Common descent
• Drake equation
• Entropy and life
• History of Earth
• List of independent discoveries#Twentieth century
• List of publications in biology#Origin of life
• Mediocrity principle
• Origin of the world's oceans
• Mimivirus
• Planetary habitability
• Rare Earth hypothesis
• Shadow biosphere
• Thermosynthesis
• Zeolite
Abiogenesis 34
Further reading
• Arrhenius, Gustaf; et al. (1997). "Entropy and Charge in Molecular Evolution—the Case of Phosphate". Journal
of Theoretical Biology 187 (4): 503–522. doi:10.1006/jtbi.1996.0385. PMID 9299295.
• Buehler, Lukas K. (2000–2005) The physico-chemical basis of life, http://www.whatislife.com/about.html
accessed 27 October 2005.
• Davies, Paul (1998). The Fifth Miracle. Penguin Science, London. ISBN 0-140-28226-2.
• De Duve, Christian (January 1996). Vital Dust: The Origin and Evolution of Life on Earth. Basic Books.
ISBN 0-465-09045-1.
• Fernando CT, Rowe, J (2007). "Natural selection in chemical evolution". Journal of Theoretical Biology 247 (1):
152–67. doi:10.1016/j.jtbi.2007.01.028. PMID 17399743.
• Hartman, Hyman (1998). "Photosynthesis and the Origin of Life". Origins of Life and Evolution of Biospheres 28
(4–6): 515–521. doi:10.1023/A:1006548904157.
• Harris, Henry (2002). Things come to life. Spontaneous generation revisited. Oxford: Oxford University Press.
ISBN 0198515383.
• Hazen, Robert M. (December 2005). Genesis: The Scientific Quest for Life's Origins [105]. Joseph Henry Press.
ISBN 0-309-09432-1.
• Gribbon, John (1998). The Case of the Missing Neutrino's and other Curious Phenomena of the Universe.
Penguin Science, London. ISBN 0-140-28734-5.
• Horgan, J (1991). "In the beginning". Scientific American 264: 100–109. (Cited on p. 108).
• Huber, C. and Wächterhäuser, G., (1998). "Peptides by activation of amino acids with CO on (Ni,Fe)S surfaces:
implications for the origin of life". Science 281 (5377): 670–672. doi:10.1126/science.281.5377.670.
PMID 9685253. (Cited on p. 108).
• Knoll, Andrew H. (2003). Life on a Young Planet: The First Three Billion Years of Evolution on Earth. Princeton
University Press.
• Luisi, Pier Luigi (2006). The Emergence of Life: From Chemical Origins to Synthetic Biology. Cambridge
University Press.
• Martin, W. and Russell M.J. (2002). "On the origins of cells: a hypothesis for the evolutionary transitions from
abiotic geochemistry to chemoautotrophic prokaryotes, and from prokaryotes to nucleated cells" [106].
Philosophical Transactions of the Royal Society: Biological sciences 358 (1429): 59–85.
doi:10.1098/rstb.2002.1183. PMID 12594918. PMC 1693102.
• Maynard Smith, John; Szathmary, Eors (2000-03-16). The Origins of Life: From the Birth of Life to the Origin of
Language. Oxford Paperbacks. ISBN 0-19-286209-X.
• Morowitz, Harold J. (1992) "Beginnings of Cellular Life: Metabolism Recapitulates Biogenesis". Yale University
Press. ISBN 0-300-05483-1
• NASA Astrobiology Institute: Earth's Early Environment and Life [107]
• NASA Specialized Center of Research and Training in Exobiology: Gustaf O. Arrhenius [108]
• Pitsch, Stefan; Krishnamurthy, Ramanarayanan; Arrhenius, Gustaf (2000). "Concentration of Simple Aldehydes
by Sulfite-Containing Double-Layer Hydroxide Minerals: Implications for Biopoesis" [109] (abstract). Helvetica
Chimica Acta 83 (9): 2398 2411. doi:10.1002/1522-2675(20000906)83:9<2398::AID-HLCA2398>3.0.CO;2-5.
• Russell MJ, Hall AJ, Cairns-Smith AG, Braterman PS (1988). "Submarine hot springs and the origin of life".
Nature 336: 117. doi:10.1038/336117a0.
• Dedicated issue of Philosophical Transactions B on Major Steps in Cell Evolution freely available. [110]
• Dedicated issue of Philosophical Transactions B on the Emergence of Life on the Early Earth freely available.
[111]
Abiogenesis 35
External links
• The Deep Hot Biosphere Theory - Thomas Gold, article from PNAS Proc. Natl. Acad. Sci. USA Vol. 89, pp.
6045-6049, July 1992 Microbiology [112]
• The Origin of Life [113] Video by John Maynard Smith
• "Harvard Team Creates the World's 1st Synthesized Cells" [114]
• Martin M Hanczyc and Jack W Szostak. Replicating vesicles as models of primitive cell growth and division.
Current Opinion in Chemical Biology 2004, 8:660–664. [115]PDF (192 KB)
• Martin A. Nowak and Hisashi Ohtsuki.Prevolutionary dynamics and the origin of evolution. Proceedings of the
National Academy of Sciences 2008 [116]
• "Exploring Life's Origins: a Virtual Exhibit" [117]
• "SELF-REPLICATION: Even peptides do it" [118] by Stuart A. Kauffman (web archive version as original page no longer
accessible)
• Origins of Life website including papers, resources, by Dr. Michael Russell at the U. of Glasgow [119]
• Possible Connections Between Interstellar Chemistry and the Origin of Life on the Earth [120]
• Scientists Find Clues That Life Began in Deep Space—NASA Astrobiology Institute [121]
• Self-organizing biochemical cycles—by Leslie Orgel [122]
• How Life Began: New Research Suggests Simple Approach [123]
• Primordial Soup's On: Scientists Repeat Evolution's Most Famous Experiment [124]–an article in Scientific
American. March 28, 2007
• Illustrations from Evolution (textbook) [125]
• An abiogenesis primer for laymen [126]
References
[1] Miller-Urey Experiment: Amino Acids & The Origins of Life on Earth (http:/ / www. juliantrubin. com/ bigten/ miller_urey_experiment.
html)
[2] Zimmer C (August 2009). "Origins. On the origin of eukaryotes". Science 325 (5941): 666–8. doi:10.1126/science.325_666.
PMID 19661396.
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[120] http:/ / pokey. arc. nasa. gov/ ~astrochm/ LifeImplications. html
[121] http:/ / nai. arc. nasa. gov/ news_stories/ news_detail. cfm?ID=207
[122] http:/ / www. pnas. org/ cgi/ content/ full/ 97/ 23/ 12503?maxtoshow=& HITS=10& hits=10& RESULTFORMAT=&
fulltext=biochemical+ cycles& searchid=1119837712082_3423& stored_search=& FIRSTINDEX=0& journalcode=pnas
[123] http:/ / www. livescience. com/ animalworld/ 060609_life_origin. html
[124] http:/ / sciam. com/ article. cfm?chanID=sa004& articleID=9952573C-E7F2-99DF-32F2928046329479
[125] http:/ / www. evolution-textbook. org/ content/ free/ figures/ ch04. html
[126] http:/ / student. science. uva. nl/ ~jckastel/ html/ abiogenesis. pdf
Miller–Urey experiment
The Miller–Urey experiment[1] (or

Urey–Miller experiment)[2] was an
experiment that simulated hypothetical
conditions thought at the time to be
present on the early Earth, and tested
for the occurrence of chemical
evolution. Specifically, the experiment
tested Alexander Oparin's and J. B. S.
Haldane's hypothesis that conditions
on the primitive Earth favored
chemical reactions that synthesized
organic compounds from inorganic
precursors. Considered to be the
classic experiment on the origin of life,
it was conducted in 1952[3] and
published in 1953 by Stanley Miller
and Harold Urey at the University of
Chicago.[4] [5] [6]
In 2008,[7] a re-analysis of Miller's The experiment
archived solutions from the original

experiments showed that 22 amino acids rather than 5 were actually created in one of the apparatus used.[8]
However, advances in science since the original experiment have shown that the conditions of the experiment do not
match that of early Earth as originally proposed.[9]
Experiment
The experiment used water (H2O), methane (CH4), ammonia (NH3), and hydrogen (H2). The chemicals were all
sealed inside a sterile array of glass tubes and flasks connected in a loop, with one flask half-full of liquid water and
another flask containing a pair of electrodes. The liquid water was heated to induce evaporation, sparks were fired
between the electrodes to simulate lightning through the atmosphere and water vapor, and then the atmosphere was
cooled again so that the water could condense and trickle back into the first flask in a continuous cycle.
At the end of one week of continuous operation, Miller and Urey observed that as much as 10–15% of the carbon
within the system was now in the form of organic compounds. Two percent of the carbon had formed amino acids
that are used to make proteins in living cells, with glycine as the most abundant. Sugars, lipids, and some of the
building blocks for nucleic acids were also formed.
In an interview, Stanley Miller stated: "Just turning on the spark in a basic pre-biotic experiment will yield 11 out of
20 amino acids."[10]
As observed in all subsequent experiments, both left-handed (L) and right-handed (D) optical isomers were created
in a racemic mixture.
The original experiment remains today under the care of Miller and Urey's former student Professor Jeffrey Bada at
the University of California, San Diego, Scripps Institution of Oceanography.[11]
Chemistry of experiment
One-step reactions among the mixture components can produce hydrogen cyanide (HCN), formaldehyde (CH2O),[12]
[13]
and other active intermediate compounds (acetylene, cyanoacetylene, etc.):
CO2 → CO + [O] (atomic oxygen)
CH4 + 2[O] → CH2O + H2O
CO + NH3 → HCN + H2O
CH4 + NH3 → HCN + 3H2 (BMA process)
The formaldehyde, ammonia, and HCN then react by Strecker synthesis to form amino acids and other biomolecules:
CH2O + HCN + NH3 → NH2-CH2-CN + H2O
NH2-CH2-CN + 2H2O → NH3 + NH2-CH2-COOH (glycine)
Furthermore, water and formaldehyde can react via Butlerov's reaction to produce various sugars like ribose.
Other experiments
This experiment inspired many others. In 1961, Joan Oró found that the nucleotide base adenine could be made from
hydrogen cyanide (HCN) and ammonia in a water solution. His experiment produced a large amount of adenine,
which molecules were formed from 5 molecules of HCN.[14] Also, many amino acids are formed from HCN and
ammonia under these conditions.[15] Experiments conducted later showed that the other RNA and DNA nucleobases
could be obtained through simulated prebiotic chemistry with a reducing atmosphere.[16]
There also had been similar electric discharge experiments related to the origin of life contemporaneous with
Miller–Urey. An article in The New York Times (March 8, 1953:E9), titled "Looking Back Two Billion Years"
describes the work of Wollman (William) M. MacNevin at The Ohio State University, before the Miller Science
paper was published in May 1953. MacNevin was passing 100,000 volt sparks through methane and water vapor and
produced "resinous solids" that were "too complex for analysis." The article describes other early earth experiments
being done by MacNevin. It is not clear if he ever published any of these results in the primary scientific literature.
K. A. Wilde submitted a paper to Science on December 15, 1952, before Miller submitted his paper to the same
journal on February 14, 1953. Wilde's paper was published on July 10, 1953.[17] Wilde used voltages up to only 600
V on a binary mixture of carbon dioxide (CO2) and water in a flow system. He observed only small amounts of
carbon dioxide reduction to carbon monoxide, and no other significant reduction products or newly formed carbon
compounds. Other researchers were studying UV-photolysis of water vapor with carbon monoxide. They have found
that various alcohols, aldehydes and organic acids were synthesized in reaction mixture [18] .
More recent experiments by chemist Jeffrey Bada at Scripps Institution of Oceanography (in La Jolla, CA) were
similar to those performed by Miller. However, Bada noted that in current models of early Earth conditions, carbon
dioxide and nitrogen (N2) create nitrites, which destroy amino acids as fast as they form. However, the early Earth
may have had significant amounts of iron and carbonate minerals able to neutralize the effects of the nitrites. When
Bada performed the Miller-type experiment with the addition of iron and carbonate minerals, the products were rich
in amino acids. This suggests the origin of significant amounts of amino acids may have occurred on Earth even with
an atmosphere containing carbon dioxide and nitrogen.[19]
Earth's early atmosphere

Some evidence suggests that Earth's original atmosphere might have contained fewer of the reducing molecules than
was thought at the time of the Miller–Urey experiment. There is abundant evidence of major volcanic eruptions 4
billion years ago, which would have released carbon dioxide, nitrogen, hydrogen sulfide (H2S), and sulfur dioxide
(SO2) into the atmosphere. Experiments using these gases in addition to the ones in the original Miller–Urey
experiment have produced more diverse molecules. The experiment created a mixture that was racemic (containing
both L and D enantiomers) and experiments since have shown that "in the lab the two versions are equally likely to
appear."[20] However, in nature, L amino acids dominate; later experiments have confirmed disproportionate
amounts of L or D oriented enantiomers are possible.[21]
Originally it was thought that the primitive secondary atmosphere contained mostly ammonia and methane.
However, it is likely that most of the atmospheric carbon was CO2 with perhaps some CO and the nitrogen mostly
N2. In practice gas mixtures containing CO, CO2, N2, etc. give much the same products as those containing CH4 and
NH3 so long as there is no O2. The hydrogen atoms come mostly from water vapor. In fact, in order to generate
aromatic amino acids under primitive earth conditions it is necessary to use less hydrogen-rich gaseous mixtures.
Most of the natural amino acids, hydroxyacids, purines, pyrimidines, and sugars have been produced in variants of
the Miller experiment.[22]
More recent results may question these conclusions. The University of Waterloo and University of Colorado
conducted simulations in 2005 that indicated that the early atmosphere of Earth could have contained up to 40
percent hydrogen—implying a possibly much more hospitable environment for the formation of prebiotic organic
molecules. The escape of hydrogen from Earth's atmosphere into space may have occurred at only one percent of the
rate previously believed based on revised estimates of the upper atmosphere's temperature.[23] One of the authors,
Owen Toon notes: "In this new scenario, organics can be produced efficiently in the early atmosphere, leading us
back to the organic-rich soup-in-the-ocean concept... I think this study makes the experiments by Miller and others
relevant again." Outgassing calculations using a chondritic model for the early earth complement the
Waterloo/Colorado results in re-establishing the importance of the Miller–Urey experiment.[24]
Conditions similar to those of the Miller–Urey experiments are present in other regions of the solar system, often
substituting ultraviolet light for lightning as the energy source for chemical reactions. The Murchison meteorite that
fell near Murchison, Victoria, Australia in 1969 was found to contain over 90 different amino acids, nineteen of
which are found in Earth life. Comets and other icy outer-solar-system bodies are thought to contain large amounts
of complex carbon compounds (such as tholins) formed by these processes, darkening surfaces of these bodies.[25]
The early Earth was bombarded heavily by comets, possibly providing a large supply of complex organic molecules
along with the water and other volatiles they contributed. This has been used to infer an origin of life outside of
Earth: the panspermia hypothesis.
Recent related studies

In recent years, studies have been made of the amino acid composition of the products of "old" areas in "old" genes,
defined as those that are found to be common to organisms from several widely separated species, assumed to share
only the last universal ancestor (LUA) of all extant species. These studies found that the products of these areas are
enriched in those amino acids that are also most readily produced in the Miller–Urey experiment. This suggests that
the original genetic code was based on a smaller number of amino acids – only those available in prebiotic nature –
than the current one.[26]
In 2008, a group of scientists examined 11 vials left over from Miller's experiments of the early 1950s. In addition to
the classic experiment, reminiscent of Charles Darwin's envisioned "warm little pond", Miller had also performed
more experiments, including one with conditions similar to those of volcanic eruptions. This experiment had a
nozzle spraying a jet of steam at the spark discharge. By using high-performance liquid chromatography and mass
spectrometry, the group found more organic molecules than Miller had. Interestingly, they found that the
volcano-like experiment had produced the most organic molecules, 22 amino acids, 5 amines and many hydroxylated
molecules, which could have been formed by hydroxyl radicals produced by the electrified steam. The group
suggested that volcanic island systems became rich in organic molecules in this way, and that the presence of
carbonyl sulfide there could have helped these molecules form peptides.[7] [27]
See also
• Abiogenesis, the study of how life on Earth emerged from inanimate organic and inorganic molecules.
External links
• A Production of Amino Acids Under Possible Primitive Earth Conditions [28] by Stanley L. Miller, Science, v.117,
May 15, 1953
• A simulation of the Miller–Urey Experiment along with a video Interview with Stanley Miller [29] by Scott Ellis
from CalSpace (UCSD)
• Origin-Of-Life Chemistry Revisited: Reanalysis of famous spark-discharge experiments reveals a richer
collection of amino acids were formed. [30]
• http://www.chem.duke.edu/~jds/cruise_chem/Exobiology/miller.html - Miller–Urey experiment explained.
References
[1] Hill HG, Nuth JA (2003). "The catalytic potential of cosmic dust: implications for prebiotic chemistry in the solar nebula and other
protoplanetary systems". Astrobiology 3 (2): 291–304. doi:10.1089/153110703769016389. PMID 14577878.
[2] Balm SP, Hare J.P., Kroto HW (1991). "The analysis of comet mass spectrometric data". Space Science Reviews 56: 185–9.
doi:10.1007/BF00178408.
[3] Bada, Jeffrey L. (2000). "Stanley Miller's 70th Birthday" (http:/ / www. issol. org/ miller/ 70thB-Day. pdf) (PDF). Origins of Life and
Evolution of the Biosphere (Netherlands: Kluwer Academic Publishers) 30: 107–12. doi:10.1023/A:1006746205180. .
[4] Miller, Stanley L. (May 1953). "Production of Amino Acids Under Possible Primitive Earth Conditions" (http:/ / www. issol. org/ miller/
miller1953. pdf) (PDF). Science 117 (3046): 528. doi:10.1126/science.117.3046.528. PMID 13056598. .
[5] Miller, Stanley L.; Harold C. Urey (July 1959). "Organic Compound Synthesis on the Primitive Earth". Science 130 (3370): 245.
doi:10.1126/science.130.3370.245. PMID 13668555. Miller states that he made "A more complete analysis of the products" in the 1953
experiment, listing additional results.
[6] A. Lazcano, J. L. Bada (June 2004). "The 1953 Stanley L. Miller Experiment: Fifty Years of Prebiotic Organic Chemistry". Origins of Life
and Evolution of Biospheres 33 (3): 235–242. doi:10.1023/A:1024807125069. PMID 14515862.
[7] Johnson AP, Cleaves HJ, Dworkin JP, Glavin DP, Lazcano A, Bada JL (October 2008). "The Miller volcanic spark discharge experiment".
Science 322 (5900): 404. doi:10.1126/science.1161527. PMID 18927386.
[8] Catherine Brahic. "Volcanic lightning may have sparked life on Earth — earth — 16 October 2008 — New Scientist Environment" (http:/ /
environment. newscientist. com/ channel/ earth/ dn14966-volcanic-lightning-may-have-sparked-life-on-earth.
html?feedId=online-news_rss20). NewScientist. . Retrieved 2008-10-17.
[9] http:/ / www. chem. duke. edu/ ~jds/ cruise_chem/ Exobiology/ miller. html
[10] "EXOBIOLOGY: An Interview with Stanley L. Miller" (http:/ / www. accessexcellence. org/ WN/ NM/ miller. php). Accessexcellence.org.
. Retrieved 2009-08-20.
[11] "A Conversation With Jeffrey L. Bada: A Marine Chemist Studies How Life Began" (http:/ / www. nytimes. com/ 2010/ 05/ 18/ science/
18conv. html). nytimes.com. 2010-05-17. .
[12] http:/ / www. oocities. com/ capecanaveral/ lab/ 2948/ orgel. html Origin of Life on Earth by Leslie E. Orgel
[13] http:/ / books. nap. edu/ openbook. php?record_id=11860& page=85 Exploring Organic Environments in the Solar System (2007)
[14] Oró J, Kimball AP (August 1961). "Synthesis of purines under possible primitive earth conditions. I. Adenine from hydrogen cyanide".
Archives of biochemistry and biophysics 94: 217–27. doi:10.1016/0003-9861(61)90033-9. PMID 13731263.
[15] Oró J, Kamat SS (April 1961). "Amino-acid synthesis from hydrogen cyanide under possible primitive earth conditions". Nature 190:
442–3. doi:10.1038/190442a0. PMID 13731262.
[16] Oró J (1967). Fox SW. ed. Origins of Prebiological Systems and of Their Molecular Matrices. New York Academic Press. pp. 137.
[17] Wilde, Kenneth A.; Bruno J. Zwolinski and Ransom B. Parlin (July 1953). "The Reaction Occurring in CO2, 2O Mixtures in a
High-Frequency Electric Arc" (http:/ / www. sciencemag. org/ cgi/ content/ citation/ 118/ 3054/ 43-a). Science 118 (3054): 43–44.
doi:10.1126/science.118.3054.43-a. PMID 13076175. . Retrieved 2008-07-09.
[18] Synthesis of organic compounds from carbon monoxide and water by UV photolysis (http:/ / www. springerlink. com/ content/
tvx0013g77u51v37/ )
[19] Fox, Douglas (2007-03-28). "Primordial Soup's On: Scientists Repeat Evolution's Most Famous Experiment" (http:/ / www. sciam. com/
article. cfm?id=primordial-soup-urey-miller-evolution-experiment-repeated). Scientific American (Scientific American Inc.). . Retrieved
2008-07-09.
[20] "Right-handed amino acids were left behind" (http:/ / www. newscientist. com/ channel/ life/ mg19025545.
200-righthanded-amino-acids-were-left-behind. html). New Scientist (Reed Business Information Ltd) (2554): pp. 18. 2006-06-02. . Retrieved
2008-07-09.
[21] Kojo, Shosuke; Hiromi Uchino, Mayu Yoshimura and Kyoko Tanaka (October 2004). "Racemic D,L-asparagine causes enantiomeric excess
of other coexisting racemic D,L-amino acids during recrystallization: a hypothesis accounting for the origin of L-amino acids in the
biosphere". Chemical Communications (19): 2146–2147. doi:10.1039/b409941a. PMID 15467844.
[22] "MICR 425: PHYSIOLOGY & BIOCHEMISTRY of MICROORGANISMS: The Origin of Life" (http:/ / www. science. siu. edu/
microbiology/ micr425/ 425Notes/ 14-OriginLife. html). SIUC / College of Science. . Retrieved 2005-12-17.
[23] "Early Earth atmosphere favorable to life: study" (http:/ / newsrelease. uwaterloo. ca/ news. php?id=4348). University of Waterloo. .
[24] Fitzpatrick, Tony (2005). "Calculations favor reducing atmosphere for early earth – Was Miller–Urey experiment correct?" (http:/ /
news-info. wustl. edu/ news/ page/ normal/ 5513. html). Washington University in St. Louis. . Retrieved 2005-12-17.
[25] Thompson WR, Murray BG, Khare BN, Sagan C (December 1987). "Coloration and darkening of methane clathrate and other ices by
charged particle irradiation: applications to the outer solar system". Journal of geophysical research 92 (A13): 14933–47.
doi:10.1029/JA092iA13p14933. PMID 11542127.
[26] Brooks D.J., Fresco J.R., Lesk A.M. & Singh M. (October 1, 2002). "Evolution of amino acid frequencies in proteins over deep time:
inferred order of introduction of amino acids into the genetic code" (http:/ / mbe. oupjournals. org/ cgi/ content/ full/ 19/ 10/ 1645). Molecular
Biology and Evolution 19 (10): 1645–55. PMID 12270892. .
[27] "'Lost' Miller–Urey Experiment Created More Of Life's Building Blocks" (http:/ / www. sciencedaily. com/ releases/ 2008/ 10/
081016141411. htm). Science Daily. October 17, 2008. . Retrieved 2008-10-18.
[28] http:/ / www. issol. org/ miller/ miller1953. pdf
[29] http:/ / millerureyexperiment. com
[30] http:/ / pubs. acs. org/ cen/ news/ 86/ i42/ 8642notw4. html
Biogenesis
Biogenesis is the process of lifeforms producing other lifeforms, e.g. a spider lays eggs, which develop into spiders.
It may also refer to biochemical processes of production in living organisms.
Generatio spontanea
The ancient Greeks believed that living things could spontaneously come into being from nonliving matter, and that
the goddess Gaia could make life arise spontaneously from stones – a process known as Generatio spontanea.
Aristotle disagreed, but he still believed that creatures could arise from dissimilar organisms or from soil. Variations
of this concept of spontaneous generation still existed as late as the 17th century, but towards the end of the 17th
century a series of observations and arguments began that eventually discredited such ideas. This advance in
scientific understanding was met with much opposition, with personal beliefs and individual prejudices often
obscuring the facts.
Francesco Redi, an Italian physician, proved as early as 1668 that higher forms of life did not originate
spontaneously, but proponents of abiogenesis claimed that this did not apply to microbes and continued to hold that
these could arise spontaneously. Attempts to disprove the spontaneous generation of life from non-life continued in
the early 1800s with observations and experiments by Franz Schulze and Theodor Schwann. In 1745 John Needham
added chicken broth to a flask and boiled it. He then let it cool and waited. Microbes grew and he proposed it as an
example of spontaneous generation. In 1768 Lazzaro Spallanzani repeated Needham's experiment, but removed all
the air from the flask. No growth occurred.[1] In 1854 Heinrich Schröder (1810 - 1885) and Theodor von Dusch, and
in 1859 Schröder alone, repeats Helmholtz filtration experiment[2] and show that living particles can be removed
from air by filtering it through cotton-wool.
Biogenesis 45
In 1864, Louis Pasteur finally announced the results of his scientific experiments. In a series of experiments similar
to those performed earlier by Needham and Spallanzani, Pasteur demonstrated that life today does not arise in areas
that have not been contaminated by existing life. Pasteur's empirical results were summarized in the phrase, Omne
vivum ex vivo, Latin for, "all life [is] from life".[3] [4]
Law of Biogenesis
The Law of Biogenesis, attributed to Louis Pasteur, states that life forms such as mice, maggots, and bacteria
produce after their own, that life does not spontaneously arise from non-life.[5] [6] Pasteur's (and others) empirical
results were summarized in the phrase, Omne vivum ex vivo, Latin for, "all life [is] from life", also known as the
"law of biogenesis". Pasteur stated: "La génération spontanée est une chimère" ("Spontaneous generation is a
dream").
See also
• Abiogenesis
• Louis Pasteur
• Aristotle
• Francesco Redi
• Franz Schulze
• Theodor Schwann
• orthogenesis
References
[1] The controversy over spontaneous generation (http:/ / www. studentsguide. in/ microbiology/ microbiology/
controversy-over-spontaneous-generation. html)
[2] McKendrick, John Gray (1899). Hermann Ludwig Ferdinand von Helmholtz (http:/ / books. google. com/ books?id=T2wPAAAAYAAJ&
lpg=PA29& ots=epwDJv1DEz& dq=helmholtz cotton putrefaction& pg=PA29#v=onepage& q& f=false). London: Fisher Unwin. pp. 162.
ISBN 9781150667695. .
[3] The microbial world: a look at things small (http:/ / www. microbiologytext. com/ index. php?module=Book& func=displayarticle&
art_id=27)
[4] Biogenesis and Abiogenesis: Critiques and Addresses (http:/ / aleph0. clarku. edu/ huxley/ CE8/ B-Ab. html)
[5] Pasteur's Papers on the Germ Theory (http:/ / biotech. law. lsu. edu/ cphl/ history/ articles/ pasteur. htm)
[6] Louis Pasteur: External links
Entropy and life 46
Entropy and life

Much writing has been devoted to entropy and life. Research concerning the relationship between the
thermodynamic quantity entropy and the evolution of life began in around the turn of the 20th century. In 1910,
American historian Henry Adams printed and distributed to university libraries and history professors the small
volume A Letter to American Teachers of History proposing a theory of history based on the second law of
thermodynamics and the principle of entropy.[1] [2] The 1944 book What is Life? by Nobel-laureate physicist Erwin
Schrödinger served largely to stimulate this research. In this book, Schrödinger states that life feeds on negative
entropy, or negentropy as it is sometimes called. Recent writings have utilized the concept of Gibbs free energy to
elaborate on this issue.
Origin
In 1863, Rudolf Clausius published his noted memoir "On the Concentration of Rays of Heat and Light, and on the
Limits of its Action" wherein he outlined a preliminary relationship, as based on his own work and that of William
Thomson, between his newly developed concept of entropy and life. Building on this, one of the first to speculate on
a possible thermodynamic perspective of evolution was the Austrian physicist Ludwig Boltzmann. In 1875, building
on the works of Clausius and Kelvin, Boltzmann reasoned:
The general struggle for existence of animate beings is not a struggle for raw materials – these, for organisms,
are air, water and soil, all abundantly available – nor for energy which exists in plenty in any body in the form
of heat, but a struggle for [negative] entropy, which becomes available through the transition of energy from
the hot sun to the cold earth.[3]
The solar system from a thermodynamic systems perspective.
Early views
In 1876, American civil engineer Richard Sears McCulloch, in his Treatise on the Mechanical Theory of Heat and
its Application to the Steam-Engine, which was an early thermodynamics textbook, states, after speaking about the
laws of the physical world, that "there are none that are established on a firmer basis than the two general
propositions of Joule and Carnot; which constitute the fundamental laws of our subject." McCulloch then goes on to
show that these two laws may be combined in a single expression as follows:
where
Entropy and life 47
= entropy
= equals a differential amount of heat passed into a thermodynamic system
= absolute temperature
McCulloch then declares that the applications of these two laws, i.e. what are currently known as the first law of
thermodynamics and the second law of thermodynamics, are innumerable. He then states:
When we reflect how generally physical phenomena are connected with thermal changes and relations, it at
once becomes obvious that there are few, if any, branches of natural science which are not more or less
dependent upon the great truths under consideration. Nor should it, therefore, be a matter of surprise that
already, in the short space of time, not yet one generation, elapsed since the mechanical theory of heat has
been freely adopted, whole branches of physical science have been revolutionized by it.
McCulloch then gives a few examples of what he calls the “more interesting examples” of the application of these
laws in extent and utility. The first example he gives, is physiology wherein he states that “the body of an animal, not
less than a steamer, or a locomotive, is truly a heat engine, and the consumption of food in the one is precisely
analogous to the burning of fuel in the other; in both, the chemical process is the same: that called combustion.” He
then incorporates a discussion of Lavoisier’s theory of respiration with cycles of digestion and excretion,
perspiration, but then contradicts Lavoisier with recent findings, such as internal heat generated by friction,
according to the new theory of heat, which, according to McCulloch, states that the “heat of the body generally and
uniformly is diffused instead of being concentrated in the chest”. McCulloch then gives an example of the second
law, where he states that friction, especially in the smaller blooded-vessels, must develop heat. Without doubt,
animal heat is thus in part produced. He then asks: “but whence the expenditure of energy causing that friction, and
which must be itself accounted for?"
To answer this question he turns to the mechanical theory of heat and goes on to loosely outline how the heart is
what he calls a “force-pump”, which receives blood and sends it to every part of the body, as discovered by William
Harvey, that “acts like the piston of an engine and is dependent upon and consequently due to the cycle of nutrition
and excretion which sustains physical or organic life.” It is likely, here, that McCulloch was modeling parts of this
argument on that of the famous Carnot cycle. In conclusion, he summarizes his first and second law argument as
such:
Everything physical being subject to the law of conservation of energy, it follows that no physiological action
can take place except with expenditure of energy derived from food; also, that an animal performing
mechanical work must from the same quantity of food generate less heat than one abstaining from exertion,
the difference being precisely the heat equivalent of that of work.
What is life?
Later, building on this premise, in the famous 1944 book What is Life?, Nobel-laureate physicist Erwin Schrödinger
theorizes that life, contrary to the general tendency dictated by the Second law of thermodynamics, decreases or
maintains its entropy by feeding on negative entropy.[4] In a note to What is Life?, however, Schrödinger explains his
usage of this term:
Let me say first, that if I had been catering for them [physicists] alone I should have let the discussion turn on
free energy instead. It is the more familiar notion in this context. But this highly technical term seemed
linguistically too near to energy for making the average reader alive to the contrast between the two things.
This is what is argued to differentiate life from other forms of matter organization. In this direction, although life's
dynamics may be argued to go against the tendency of second law, which states that the entropy of an isolated
system tends to increase, it does not in any way conflict or invalidate this law, because the principle that entropy can
only increase or remain constant applies only to a closed system which is adiabatically isolated, meaning no heat can
enter or leave. Whenever a system can exchange either heat or matter with its environment, an entropy decrease of
Entropy and life 48
that system is entirely compatible with the second law.[5]

In 1964, James Lovelock was among a group of scientists who were requested by NASA to make a theoretical life
detection system to look for life on Mars during the upcoming space mission. When thinking about this problem,
Lovelock wondered “how can we be sure that Martian life, if any, will reveal itself to tests based on Earth’s
lifestyle?” [6] To Lovelock, the basic question was “What is life, and how should it be recognized?” When speaking
about this puzzling issue with some of his colleagues at the Jet Propulsion Laboratory, he was asked, well what
would you do to look for life on Mars? To this Lovelock replied:
I’d look for an entropy reduction, since this must be a general characteristic of life.
Thus, according to Lovelock, to find signs of life, one must look for a “reduction or a reversal of entropy.”
Gibbs free energy and biological evolution

In recent years, the thermodynamic interpretation of evolution in relation to entropy has begun to utilize the concept
of the Gibbs free energy, rather than entropy. This is because biological processes on earth take place at roughly
constant temperature and pressure, a situation in which the Gibbs free energy is an especially useful way to express
the second law of thermodynamics. The Gibbs free energy is given by:
The minimization of the Gibbs free energy is a form of the principle of minimum energy, which follows from the
entropy maximization principle for closed systems. Moreover, the Gibbs free energy equation, in modified form, can
be utilized for open systems when chemical potential terms are included in the energy balance equation. In a popular
1982 textbook Principles of Biochemistry by noted American biochemist Albert Lehninger, it is argued that the order
produced within cells as they grow and divide is more than compensated for by the disorder they create in their
surroundings in the course of growth and division. In short, according to Lehninger, "living organisms preserve their
internal order by taking from their surroundings free energy, in the form of nutrients or sunlight, and returning to
their surroundings an equal amount of energy as heat and entropy."[7]
Similarly, according to the chemist John Avery, from his recent 2003 book Information Theory and Evolution, we
find a presentation in which the phenomenon of life, including its origin and evolution, as well as human cultural
evolution, has its basis in the background of thermodynamics, statistical mechanics, and information theory. The
(apparent) paradox between the second law of thermodynamics and the high degree of order and complexity
produced by living systems, according to Avery, has its resolution "in the information content of the Gibbs free
energy that enters the biosphere from outside sources."[8] The process of natural selection responsible for such local
increase in order may be mathematically derived directly from the expression of the second law equation for
connected non-equilibrium open systems [9] .
Entropy and the origin of life

The second law of thermodynamics applied on the origin of life is a far more complicated issue than the further
development of life, since there is no "standard model" model of how the first biological lifeforms emerged; only a
number of competing hypotheses. The problem is discussed within the area of Abiogenesis, implying gradual
pre-Darwinian chemical evolution. In 1924, Alexander Oparin suggested that sufficient energy was provided in a
primordial soup. The Belgian scientist Ilya Prigogine was awarded with a Nobel prize for an analysis in this area. A
related topic is the probability that life would emerge, which has been discussed in several studies, for example by
Russell Doolittle.[10]
Entropy and life 49
See also
• Complexity theory
• Dissipative system
• Entropy (order and disorder)
Further reading
• Schneider, E. and Sagan, D. (2005). Into the Cool: Energy Flow, Thermodynamics, and Life. University of
Chicago Press, Chicago.
• La Cerra, P. (2003). The First Law of Psychology is the Second Law of Thermodynamics: The Energetic
Evolutionary Model of the Mind and the Generation of Human Psychological Phenomena. Human Nature
Review, Volume 3: 440-447. Full text [11]
External links
• Life on Earth - Flow of Energy and Entropy [12] - Marek Roland-Mieszkowski, M.Sc., Ph. D.,
• Thermodynamic Evolution of the Universe [13]
References
[1] Adams, Henry. (1986). History of the United States of America During the Administration of Thomas Jefferson (pg. 1299). Library of
America.
[2] Adams, Henry. (1910). A Letter to American Teachers of History. Google Books (http:/ / books. google. com/ books?id=gaLdOOzuiKAC&
pg=PA1& dq=A+ Letter+ to+ American+ Teachers+ of+ History#PPA10,M1), Scanned PDF (http:/ / ia311517. us. archive. org/ 0/ items/
alettertoamerica00adamuoft/ alettertoamerica00adamuoft. pdf). Washington.
[3] Boltzmann, Ludwig (1974). The second law of thermodynamics (Theoretical physics and philosophical problems). Springer-Verlag New
York, LLC. ISBN 9789027702500.
[4] Schrödinger, Erwin (1944). What is Life - the Physical Aspect of the Living Cell. Cambridge University Press. ISBN 0-521-42708-8.
[5] The common justification for this argument, for example, according to renowned chemical engineer Kenneth Denbigh, from his 1955 book
The Principles of Chemical Equilibrium, is that "living organisms are open to their environment and can build up at the expense of foodstuffs
which they take in and degrade."
[6] Lovelock, James (1979). GAIA - A New Look at Life on Earth. Oxford University Press. ISBN 0-19-286218-9.
[7] Lehninger, Albert (1993). Principles of Biochemistry, 2nd Ed.. Worth Publishers. ISBN 0-87901-711-2.
[8] Avery, John (2003). Information Theory and Evolution. World Scientific. ISBN 981-238-399-9.
[9] Kaila, V. R. and Annila, A. (8 November 2008). "Natural selection for least action". Proceedings of the Royal Society A 464 (2099):
3055–3070. doi:10.1098/rspa.2008.0178.
[10] Russell Doolittle, "The Probability and Origin of Life" in Scientists Confront Creationism (1984) Ed. Laurie R. Godfrey, p. 85
[11] http:/ / human-nature. com/ nibbs/ 03/ lacerra. pdf
[12] http:/ / www. digital-recordings. com/ publ/ publife. html
[13] http:/ / pi. physik. uni-bonn. de/ ~cristinz/ thesis/ t/ node7. html
Mimivirus 50
Mimivirus
Mimivirus
Virus classification
Group: Group I (dsDNA)
(unranked): Nucleocytoplasmic large DNA viruses
Family: Mimiviridae
Genus: Mimivirus
Species: Acanthamoeba polyphaga mimivirus
Mimivirus is a viral genus containing a single identified species named Acanthamoeba polyphaga mimivirus
(APMV), or is a group of (designated usually MimiN) phylogenetically related large viruses[1] . In colloquial speech,
APMV is more commonly referred to as just “mimivirus”. It has the largest capsid diameter of all known viruses, as
well as a large and complex genome compared with other viruses. Though knowledge of the virus is relatively
limited, the discovery of the virus excited many people due to the implications of its complex nature, with people
hailing it as everything from a new domain of life to a missing link between viruses and bacteria.
Discovery
APMV was discovered serendipitously in 1992 within the amoeba Acanthamoeba polyphaga, after which it is
named, during research into Legionellosis. The virus was observed in a gram stain and mistakenly thought to be a
gram-positive bacterium. As a consequence it was named "Bradfordcoccus", after the district the amoeba was
sourced from in Bradford, England. In 2003, researchers at the Université de la Méditerranée in Marseille, France
published a paper in Science identifying the micro-organism as a virus.[2]
Mimivirus may be a causative agent of some forms of pneumonia; this is based mainly on indirect evidence in the
form of antibodies to the virus discovered in pneumonia patients.[3] Although the classification of mimivirus as a
pathogen is tentative, evidence is accumulating that it can cause viral pneumonia.[4]
Classification
It has not been placed into a viral family by the International Committee on Taxonomy of Viruses but more members
of the proposed family Mimiviridae are thought to exist based on metagenomic data.[5] It has however, been placed
into Group I of the Baltimore classification system.
Whilst not strictly a method of classification, Mimivirus joins a group of large viruses known as nucleocytoplasmic
large DNA viruses (NCLDV). They are all large viruses which share both molecular characteristics and large
genomes. The mimivirus genome also possesses 21 genes encoding homologs to proteins which are seen to be highly
conserved in the majority of NCLDVs, and further work suggests that mimivirus is an early divergent of the general
NCLDV group.[2]
Mimivirus 51
Structure
The mimivirus is larger than all previously discovered viruses, with a capsid diameter of 400 nm. Protein filaments
measuring 100 nm project from the surface of the capsid, bringing the total length of the virus up to 600 nm.
Variation in scientific literature renders these figures as highly approximate, with the "size" of the virion being
casually listed as anywhere between 400 nm and 800 nm, depending on whether total length or capsid diameter is
actually quoted. The capsid appears hexagonal under an electron microscope, therefore the capsid symmetry is
icosahedral.[6] It does not appear to possess an outer viral envelope, suggesting that the virus does not exit the host
cell by exocytosis.[7]
The same team that discovered the mimivirus later discovered a slightly larger virus, dubbed the mamavirus, and the
Sputnik virophage that infects it[8] .
Mimivirus shares several morphological characteristics with all members of the NCLDV group of viruses. As an
internal lipid layer surrounding the central core is present in all other NCLDV viruses, it has been suggested by M.
Suzan-Monti et al. that this may also be present in mimivirus. The condensed central core of the virion appears as a
dark region under the electron microscope. The large genome of the virus resides within this area.
Several mRNA transcripts can be recovered from purified virions. Like other NCLDVs, transcripts for DNA
polymerase, a capsid protein and a TFII-like transcription factor were found. However, three distinct aminoacyl
tRNA synthetase enzyme transcripts and four unknown mRNA molecules specific to mimivirus were also found.
These pre-packaged transcripts can be translated without viral gene expression and are likely to be necessary to
Mimivirus for replication. Other DNA viruses, such as the Human cytomegalovirus and Herpes simplex virus type-1,
also feature pre-packaged mRNA transcripts.[7]
Genome
The mimivirus genome is a linear, double-stranded molecule of DNA with 1,181,404 base pairs in length.[9] This
makes it the largest viral genome in scientific knowledge, outstripping the next-largest virus genome of the myovirus
Bacillus phage G by a little over double.[10] In addition, it is larger than at least 30 cellular clades.[11]
In addition to the large size of the genome, mimivirus possesses an estimated 911 protein-coding genes, far
exceeding the minimum 4 genes required for viruses to exist (c.f. MS2 and Qβ viruses).[12] Analysis of its genome
revealed the presence of genes not seen in any other viruses, including aminoacyl tRNA synthetases, and other genes
previously thought only to be encoded by cellular organisms. Like other large DNA viruses, mimivirus contains
several genes for sugar, lipid and amino acid metabolism, as well as some metabolic genes not found in any other
virus.[7] Roughly 90% of the genome was of coding capacity, with the other 10% being “junk DNA”.
Replication
The stages of mimivirus replication are not well known, but as a minimum it is known that mimivirus attaches to a
chemical receptor on the surface of an amoeba cell and is taken into the cell. Once inside, an eclipse phase begins, in
which the virus disappears and all appears normal within the cell. After about four hours small accumulations can be
seen in areas of the cell. Eight hours after infection many mimivirus virions are clearly visible within the cell. The
cell cytoplasm continues to fill with newly synthesised virions and about 24 hours after initial infection the cell
likely bursts open to release the new mimivirus virions.[7]
Little is known about the details of this replication cycle, most obviously attachment to the cell surface and entry,
viral core release, DNA replication, transcription, translation, assembly and release of progeny virions. However,
scientists have established the general overview given above using electron micrographs of infected cells. These
micrographs show mimivirus capsid assembly in the nucleus, acquisition of an inner lipid membrane via budding
from the nucleus, and particles similar to those found in many other viruses, including all NCLDV members. These
particles are known in other viruses as viral factories and allow efficient viral assembly by modifying large areas of
Mimivirus 52
the host cell.
Implications for defining "life"

Mimivirus possesses many characteristics which place it at the boundary of living and non-living. It is as large as
several bacterial species, such as Rickettsia conorii and Tropheryma whipplei, possesses a genome of comparable
size to several bacteria, including those above, and codes for products previously not thought to be encoded by
viruses. In addition, mimivirus possesses genes coding for nucleotide and amino acid synthesis, which even some
small obligate intracellular bacteria lack. This means that unlike these bacteria, mimivirus is not dependent on the
host cell genome for coding the metabolic pathways for these products. They do however, lack genes for ribosomal
proteins, making mimivirus dependent on a host cell for protein translation and energy metabolism. These factors
combined have thrown scientists into debate over whether mimivirus is a distinct form of life, comparable on a
domain scale to Eukarya, Archaea and Bacteria. Nevertheless, mimivirus does not exhibit the following
characteristics, all of which are part of many conventional definitions of life: homeostasis, response to stimuli,
growth in the normal sense of the term (instead replicating via self-assembly of individual components) or
undergoing cellular division.
Because its lineage is very old and could have emerged prior to cellular organisms, mimivirus has added to the
debate over the origins of life. Some genes unique to mimivirus, including those coding for the capsid, have been
conserved in a variety of viruses which infect organisms from all domains - Eukarya, Archaea and Bacteria. This has
been used to suggest that mimivirus is related to a type of DNA virus that emerged before cellular organisms and
played a key role in the development of all life on Earth.[13] An alternative hypothesis is that there were three distinct
types of DNA viruses that were involved in generating the three known domains of life.[14]
References in popular culture

A "pseudo-mimi" drove much of the plot of Vernor Vinge's novel Rainbows End. In the book, it was a tailored virus
used for highly effective and subtle mind control. The virus's relatively large genetic capacity was central to this
idea.
Mentioned in episode 3, season 4 of the Canadian television show ReGenesis.
See also
• Mycoplasma genitalium – one of the smallest bacteria
• Pelagibacter ubique – possesses one of the smallest bacterial genomes
• Nanoarchaeum equitans – smallest known independent cell, one of the smallest known genomes (the smallest
published genome belongs to Carsonella rudii )
• Nanobacterium
• Nanobe
• Marseillevirus another giant virus
• Parvovirus – smallest known viruses
• Virophage – mimivirus is so large that it has its own parasite species.
Mimivirus 53
Further reading
• Raoult, D.; et al. (2004). "The 1.2-megabase genome sequence of Mimivirus". Science 306 (5700): 1344–1350.
doi:10.1126/science.1101485. PMID 15486256.
• Ghedin, Elodie; Claverie, J. M. (2005). "Mimivirus relatives in the Sargasso sea" [15]. Virology Journal 2: 62.
doi:10.1186/1743-422X-2-62. PMID 16105173. PMC 1215527.
• Peplow, Mark, 2004, "Giant virus qualifies as 'living organism' [16]," News@Nature, doi:10.1038/
• "Mimivirus: discovery of a giant virus" [17]. Press Release. Paris: Centre national de la recherche scientifique.
2003-03-28.
• New Scientist, Issue 2544, 25 March 2006.
• GiantVirus.org [18]
• Highfield, Roger (2004-10-15). "The Bradford bug that may be a new life form" [19]. Daily Telegraph.
• "Scientists investigate structural details of the largest known virus" [20]. Science News. 2009-04-28.
• Keim, Brandon (2009-05-05). "Viral Missing Link Caught on Film" [21]. Wired Science.
External links
• Viralzone: Mimiviridae [22]
• International Committee on Taxonomy of Viruses (ICTV) picture gallery [23] - images of mimivirus
References
[1] Ghedin, E.; Claverie, J. (Aug 2005). "Mimivirus relatives in the Sargasso sea" (http:/ / www. virologyj. com/ content/ 2/ / 62) (Free full text).
Virology journal 2: 62. doi:10.1186/1743-422X-2-62. PMID 16105173. PMC 1215527. .
[2] La Scola B, Audic S, Robert C, Jungang L, de Lamballerie X, Drancourt M, Birtles R, Claverie JM, Raoult D. (2003). "A giant virus in
amoebae". Science 299 (5615): 2033. doi:10.1126/science.1081867. PMID 12663918.
[3] La Scola B, Marrie T, Auffray J, Raoult D (2005). "Mimivirus in pneumonia patients" (http:/ / www. cdc. gov/ ncidod/ EID/ vol11no03/
04-0538. htm). Emerg Infect Dis 11 (3): 449–52. PMID 15757563. .
[4] Raoult D, Renesto P, Brouqui P (2006). "Laboratory infection of a technician by mimivirus" (http:/ / www. annals. org/ cgi/ reprint/ 144/ 9/
702-b. pdf). Ann Intern Med 144 (9): 702–3. PMID 16670147. .
[5] "ICTV entry on Mimivirus" (http:/ / www. ncbi. nlm. nih. gov/ ICTVdb/ Ictv/ fs_mimiv. htm). .
[6] Xiao C, Kuznetsov YG, Sun S, Hafenstein SL, Kostyuchenko VA, Chipman PR, Suzan-Monti M, Raoult D, McPherson A, Rossmann MG
(2009-04-28). "Structural studies of the giant mimivirus" (http:/ / www. pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez&
artid=2671561). PLoS Biol. 7 (4): e92. doi:10.1371/journal.pbio.1000092. PMID 19402750. PMC 2671561.
[7] Suzan-Monti M, La Scola B, Raoult D (2006). "Genomic and evolutionary aspects of Mimivirus". Virus Research 117 (1): 145–155.
doi:10.1016/j.virusres.2005.07.011. PMID 16181700.
[8] Nature: "'Virophage' suggests viruses are alive" (http:/ / www. nature. com/ nature/ journal/ v454/ n7205/ full/ 454677a. html) by Helen
Pearson 2008
[9] "Acanthamoeba polyphaga mimivirus, complete genome" (http:/ / www. ncbi. nlm. nih. gov/ sites/ entrez?db=genome& cmd=Retrieve&
dopt=Overview& list_uids=18057). NCBI. .
[10] "Top 100 largest viral genome sequences" (http:/ / www. giantvirus. org/ top. html). GiantVirus.org. .
[11] Claverie, Jean-Michel; et al. (2006). "Mimivirus and the emerging concept of ‘giant’ virus". Virus Research 117 (1): 133–144.
doi:10.1016/j.virusres.2006.01.008. PMID 16469402.
[12] Prescott, Lansing M. (1993). 2nd edition. ed. Microbiology. Dubuque, IA: Wm. C. Brown Publishers. ISBN 0697013723.
[13] Siebert, Charles (2006-03-15). "Unintelligent Design" (http:/ / discovermagazine. com/ 2006/ mar/ unintelligent-design). Discover
Magazine. .
[14] Forterre, Patrick (2006). "Three RNA cells for ribosomal lineages and three DNA viruses to replicate their genomes: A hypothesis for the
origin of cellular domain" (http:/ / www. pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez& artid=1450140). PNAS 106 (10):
3669–3674. doi:10.1073/pnas.0510333103. PMID 16505372. PMC 1450140.
[15] http:/ / www. pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez& artid=1215527
[16] http:/ / www. nature. com/ news/ 2004/ 041011/ full/ 041011-14. html
[17] http:/ / www. cnrs. fr/ cw/ en/ pres/ compress/ mimivirus. htm
[18] http:/ / www. giantvirus. org
[19] http:/ / www. telegraph. co. uk/ news/ main. jhtml?xml=/ news/ 2004/ 10/ 15/ nbug15. xml& sSheet=/ news/ 2004/ 10/ 15/ ixhome. html
[20] http:/ / www. sciencenews. org/ view/ generic/ id/ 43277/ title/ Mimivirus_up_close
[21] http:/ / www. wired. com/ wiredscience/ 2009/ 05/ mimivirus/
Mimivirus 54
[22] http:/ / www. expasy. org/ viralzone/ all_by_species/ 17. html

[23] http:/ / www. ncbi. nlm. nih. gov/ ICTVdb/ WIntkey/ Images/ em_mimiv. htm
Rare Earth hypothesis

In planetary astronomy and astrobiology, the Rare Earth
hypothesis argues that the emergence of complex multicellular
life (metazoa) on Earth required an improbable combination of
astrophysical and geological events and circumstances. The term
"Rare Earth" comes from Rare Earth: Why Complex Life Is
Uncommon in the Universe (2000), a book by Peter Ward, a
geologist and paleontologist, and Donald E. Brownlee, an
astronomer and astrobiologist. Their book is the source for much
of this article.
The rare earth hypothesis is the contrary of the principle of

mediocrity (also called the Copernican principle), advocated by
Carl Sagan and Frank Drake, among others.[1] The principle of
mediocrity concludes that the Earth is a typical rocky planet in a Are planets that support life, such as Earth, rare?
typical planetary system, located in an unexceptional region of a
common barred-spiral galaxy. Hence it is probable that the universe teems with complex life. Ward and Brownlee
argue to the contrary: planets, planetary systems, and galactic regions that are as friendly to complex life as are the
Earth, the solar system, and our region of the Milky Way are very rare.
By concluding that complex life is uncommon, the Rare Earth hypothesis is a possible solution to the Fermi paradox:
"If extraterrestrial aliens are common, why aren't they obvious?"[2]
Why complex life may be very rare

The Rare Earth hypothesis argues that the emergence of complex life requires a host of fortuitous circumstances. A
number of such circumstances are set out below under the following headings: galactic habitable zone, a central star
and planetary system having the requisite character, the circumstellar habitable zone, the size of the planet, the
advantage of a large satellite, conditions needed to assure the planet has a magnetosphere and plate tectonics, the
chemistry of the lithosphere, atmosphere, and oceans, the role of "evolutionary pumps" such as massive glaciation
and rare bolide impacts, and whatever led to the still mysterious Cambrian explosion of animal phyla. The
emergence of intelligent life may have required yet other rare events.
In order for a small rocky planet to support complex life, Ward and Brownlee argue, the values of several variables
must fall within narrow ranges. The universe is so vast that it could contain many Earth-like planets. But if such
planets exist, they are likely to be separated from each other by many thousands of light years. Such distances may
preclude communication among any intelligent species evolving on such planets, which would solve the Fermi
paradox.
The galactic habitable zone

Rare Earth suggests that much of the known
universe, including large parts of our galaxy,
cannot support complex life; Ward and
Brownlee refer to such regions as "dead
zones." Those parts of a galaxy where
complex life is possible make up the galactic
habitable zone. This zone is primarily a
function of distance from the galactic center.
As that distance increases:
1. The metallicity of stars declines, and In an area lacking metals or an area near the center with high radiation, a planet
metals (which in astronomy means all would not be able to sustain life (galaxy shown is NGC 7331, often referred to as
[3]
elements other than hydrogen and helium) the "twin" of the Milky Way )
are necessary to the formation of

terrestrial planets.
2. The X-ray and gamma ray radiation from the black hole at the galactic center, and from nearby neutron stars,
becomes less intense. Radiation of this nature is considered dangerous to complex life, hence the Rare Earth
hypothesis predicts that the early universe, and likewise regions in the galaxy at present where the stellar density
is high and supernovae common, will be unfit for the development of complex life.[4]
3. Gravitational perturbation of planets and planetesimals by nearby stars becomes less likely as the density of stars
decreases. Hence the further a planet lies from the galactic center, the less likely it is to be struck by a large
bolide. A sufficiently large impact may extinguish all complex life on a planet.
(1) rules out the outer reaches of a galaxy; (2) and (3) rule out galactic inner regions, globular clusters, and the spiral
arms of spiral galaxies. These arms are not physical objects, but regions of a galaxy characterized by a higher rate of
star formation, moving very slowly through the galaxy in a wave-like manner. As one moves from the center of a
galaxy to its furthest extremity, the ability to support life rises then falls. Hence the galactic habitable zone may be
ring-shaped, sandwiched between its uninhabitable center and outer reaches.
While a planetary system may enjoy a location favorable to complex life, it must also maintain that location for a
span of time sufficiently long for complex life to evolve. Hence a central star with a galactic orbit that steers clear of
galactic regions where radiation levels are high, such as the galactic center and the spiral arms, would appear most
favourable. If the central star's galactic orbit is eccentric (elliptic or hyperbolic), it will pass through some spiral
arms, but if the orbit is a near perfect circle and the orbital velocity equals the "rotational" velocity of the spiral arms,
the star will drift into a spiral arm region only gradually, if at all. Therefore Rare Earth proponents conclude that a
life-bearing star must have a galactic orbit that is nearly circular about the center of its galaxy. The required
synchronization of the orbital velocity of a central star with the wave velocity of the spiral arms can occur only
within a fairly narrow range of distances from the galactic center. This region is termed the "galactic habitable zone".
Lineweaver et al.[5] calculate that the galactic habitable zone is a ring 7 to 9 kiloparsecs in diameter, that includes no
more than 10% of the stars in the Milky Way.[6] Based on conservative estimates of the total number of stars in the
galaxy, this could represent something like 20 to 40 billion stars. Gonzalez et al.[7] would halve these numbers; he
estimates that at most 5% of stars in the Milky Way fall in the galactic habitable zone.
The orbit of the Sun around the center of the Milky Way is indeed almost perfectly circular, with a period of 226 Ma
(1 Ma = 1 million years), one closely matching the rotational period of the galaxy. Karen Masters calculated that the
orbit of the Sun takes it through a major spiral arm approximately every 100 million years. In contrast, the Rare
Earth hypothesis predicts that the Sun, since its formation, should have passed through no spiral arm at all.[8]
However, some researchers have suggested that several mass extinctions do correspond with previous crossings of
the spiral arms.[9]
A central star of the right character

The terrestrial example suggests complex life requires water in the liquid state and its planet must therefore be at an
appropriate distance. This is the core of the notion of the habitable zone or Goldilocks Principle [10] . The habitable
zone forms a ring around the central star. If a planet orbits its sun too closely or too far away, the surface temperature
is incompatible with water being liquid (though sub-surface water, as suggested for Europa, Enceladus, and Ceres,
may be possible at varying locations[11] ). Kasting et al. (1993) estimate that the habitable zone for the Sun ranges
from 0.95 to 1.15 astronomical units.[12]
The habitable zone varies with the type and
age of the central star. The habitable zone for
a main sequence star very gradually moves
out over time until the star becomes a white
dwarf, at which time the habitable zone
vanishes. The habitable zone is closely
connected to the greenhouse warming
afforded by atmospheric carbon dioxide
(CO2) or other greenhouse gases. Even
though the Earth's atmosphere contains only
387 parts per million of CO2, that trace
amount suffices to raise the average surface A Habitable zone chart showing where Earth-like life could exist (however,
temperature of the Earth by about 40°C from microbial life could exist outside the habitable zone in the image)
what it would otherwise be [13] .
It is then presumed a star needs to have rocky planets within its habitable zone. While the habitable zone of hot stars
such as Sirius or Vega is wide, there are two problems:
1. Given that rocky planets were (at the time Rare Earth was written) thought to form closer to their central stars,
the planet likely forms too close to the star to lie within the habitable zone. This does not rule out life on a moon
of a gas giant. Hot stars also emit much more ultraviolet radiation, which will ionize any planetary atmosphere.
2. Hot stars, as mentioned above, have short lives, becoming red giants in as little as 1 Ga. This may not allow
enough time for advanced life to evolve.
These considerations rule out the massive and powerful stars of type F6 to O (see stellar classification) as homes to
evolved metazoan life.
Small red dwarf stars, on the other hand, have habitable zones with a small radius. This proximity causes one face of
the planet to constantly face the star, and the other to always remain dark, a situation known as tidal lock. Tidal
locking of a planetary hemisphere to its primary will cause one side of a planet to be extremely hot, while the other
will be extremely cold. Planets within a habitable zone with a small radius are also at increased risk of solar flares
(see Aurelia), which would tend to ionize the atmosphere and are otherwise inimical to complex life. Rare Earth
proponents argue that this rules out the possibility of life in such systems, though some exobiologists have suggested
that habitability may exist under the right circumstances. This is a central point of contention for the theory, since
these late-K and M category stars make up about 82% of all hydrogen-burning stars.[14]
Rare Earth proponents argue that the stellar type of central stars that are "just right" ranges from F7 to K1. Such stars
are not common: G type stars such as the Sun (between the hotter F and cooler K) comprise only 9%[14] of the
hydrogen-burning stars in the Milky Way.
Aged stars, such as red giants and white dwarfs, are also
unlikely to support life. Red giants are common in globular
clusters and elliptical galaxies. White dwarfs are mostly dying
stars that have already gone through their red giant phase. The
diameter of a red giant has substantially increased from its
youth. If a planet was in the habitable zone during a star's
youth and middle age, it will be fried when its parent star
becomes a red giant (though theoretically planets at a much
greater distance may become habitable).
The energy output of a star over its lifespan should only change
very gradually; variable stars such as Cepheid variables, for
instance, are highly unlikely to support life. If the central star's
energy output suddenly decreases, even for a relatively short
while, the planet's water may freeze. Conversely, if the central
Globular clusters are unlikely to support life
star's energy output significantly increases, the oceans may
evaporate, resulting in a greenhouse effect; this may preclude
the oceans from reforming.
There is no known way to achieve life without complex chemistry, and such chemistry requires metals, namely
elements other than hydrogen, helium, and lithium. This suggests a condition for life is a solar system rich in metals.
The only known mechanism for creating and dispersing metals is a supernova explosion. The presence of metals in
stars is revealed by their absorption spectrum, and studies of stellar spectra reveal that many, perhaps most, stars are
poor in metals. Low metallicity characterizes the early universe, globular clusters and other stars formed when the
universe was young, stars in most galaxies other than large spirals, and stars in the outer regions of all galaxies. Thus
metal-rich central stars capable of supporting complex life are believed most common in the quiet suburbs of the
larger spiral galaxies, regions hospitable to complex life for another reason, namely the absence of high radiation.[15]
Planetary system
A gas cloud capable of giving birth to a star can also give rise to gas giant low metallicity (Jovian) planets like
Jupiter and Saturn. But Jovian planets have no hard surface of the kind believed necessary for complex life (their
satellites may have hard surfaces, though). The Ward and Brownlee argument holds that a planetary system capable
of sustaining complex life must be structured more or less like our solar system, with small and rocky inner planets,
and Jovian outer ones. Recent research calls this line of argument into question, however.
Uncertainty over Jupiter
At the time of Ward and Brownlee's book, gas giants

were thought to help support life by keeping asteroids
away from the life-bearing planet. Hence a gas giant
was thought to protect the inner rocky planets from
asteroid bombardment. However, recent computer
simulations on the matter suggest that the situation is
more complex than this; Jupiter appears to cause three
times as many asteroid impacts as it prevents, while
replacing Jupiter with a Saturn-sized body would
roughly double the bombardment rate compared to
Jupiter.[17]
Disruption of orbit
A gas giant must not be too close to a body upon which

life is developing, unless that body is one of its moons.
Close placement of gas giant(s) could disrupt the orbit Jupiter is the fifth planet from the Sun and the largest planet within
[16]
of a potential life-bearing planet, either directly or by the Solar System.
drifting into the habitable zone.

Newtonian dynamics can produce chaotic planetary orbits, especially in a system having large planets at high orbital
eccentricity.[18]
The need for stable orbits rules out stars with systems of planets that contain large planets with orbits close to the
host star (called "hot Jupiters"). It is believed that hot Jupiters formed much further from their parent stars than they
are now, and have migrated inwards to their current orbits. In the process, they would have catastrophically disrupted
the orbits of any planets in the habitable zone.[19]
Size of planet
A planet that is too small cannot hold much of an atmosphere. Hence the surface temperature becomes more variable
and the average temperature drops. Substantial and long-lasting oceans become impossible. A small planet will also
tend to have a rough surface, with large mountains and deep canyons. The core will cool faster, and plate tectonics
will either not last as long as they would on a larger planet or may not occur at all.[20]
Small rocky planets like Earth may be common according to astronomer Michael Meyer of the University of
Arizona.
Our observations suggest that between 20% and 60% of Sun-like stars have evidence for the formation of
rocky planets not unlike the processes we think led to planet Earth. That is very exciting.
—Michael Meyer, [21]
Meyer’s team found cosmic dust near recently-formed sun-like stars and sees this as a byproduct of the formation of
rocky planets.
Large moon
The Moon is unusual because the other rocky planets in the Solar System either have no satellites (Mercury and
Venus), or have tiny satellites that are likely captured asteroids (Mars).
The giant impact theory hypothesizes that the Moon resulted from the impact of a Mars-sized body, Theia, with the
very young Earth. This giant impact also gave the Earth its axis tilt and velocity of rotation [22] . Rapid rotation
reduces the daily variation in temperature and makes photosynthesis viable. The Rare Earth hypothesis further
argues that the axis tilt cannot be too large or too small (relative to the orbital plane). A planet with a large tilt will
experience extreme seasonal variations in climate, unfriendly to complex life. A planet with little or no tilt will lack
the stimulus to evolution that climate variation provides. In this view, the Earth's tilt is "just right". The gravity of a
large satellite also stabilises the planet's tilt; without this effect the variation in tilt would be chaotic, probably
making complex life forms on land impossible.[23]
If the Earth had no Moon, the ocean tides resulting solely from the Sun's gravity would be very modest. A large
satellite gives rise to tidal pools, which may be essential for the formation of complex life, though this is far from
certain.[24]
A large satellite also increases the likelihood of plate tectonics through the effect of tidal forces on the planet's crust.
The impact that formed the Moon may also have initiated plate tectonics, without which the continental crust would
cover the entire planet, leaving no room for oceanic crust. It is possible that the large scale mantle convection needed
to drive plate tectonics could not have emerged in the absence of crustal inhomogeneity. However, there is strong
evidence that plate tectonics existed on Mars in the past, without such a mechanism to initiate it.[25]
If a giant impact is the only way for a rocky inner planet to acquire a large satellite, any planet in the circumstellar
habitable zone will need to form as a double planet in order that there be an impacting object sufficiently massive to
give rise in due course to a large satellite. An impacting object of this nature is not necessarily improbable. Recent
work by Edward Belbruno and J. Richard Gott of Princeton University suggests that a suitable impacting body could
form in a planet's trojan points (L4 or L5 Lagrangian point).[26]
Plate tectonics
A planet will not experience plate tectonics unless its chemical composition allows it. The only known long lasting
source of the required heat is radioactive decay occurring deep in the planet's interior. Continents must also be made
up of less dense granitic rocks that "float" on underlying denser basaltic rock. Taylor [27] emphasizes that subduction
zones (an essential part of plate tectonics) require the lubricating action of ample water; on Earth, such zones exist
only at the bottom of oceans.
Rare Earth equation

The following discussion is adapted from Cramer.[28] The Rare Earth equation is Ward and Brownlee's riposte to the
Drake equation. It calculates , the number of Earth-like planets in the Milky Way having complex life forms, as:
[29]
where:
• N* is the number of stars in the Milky Way. This number is not well-estimated, because the Milky Way's mass is
not well estimated. Moreover, there is little information about the number of very small stars. N* is at least 100
billion, and may be as high as 500 billion, if there are many low visibility stars.
• is the average number of planets in a star's habitable zone. This zone is fairly narrow, because constrained by
the requirement that the average planetary temperature be consistent with water remaining liquid throughout the
time required for complex life to evolve. Thus = 1 is a likely upper bound.
We assume . The Rare Earth hypothesis can then be viewed as asserting that the product of the
other nine Rare Earth equation factors listed below, which are all fractions, is no greater than 10−10 and could
plausibly be as small as 10−12. In the latter case, could be as small as 0 or 1. Ward and Brownlee do not actually
calculate the value of , because the numerical values of quite a few of the factors below can only be conjectured.
They cannot be estimated simply because we have but one data point: the Earth, a rocky planet orbiting a G2 star in a
quiet suburb of a large barred spiral galaxy, and the home of the only intelligent species we know, namely ourselves.
• is the fraction of stars in the galactic habitable zone (Ward, Brownlee, and Gonzalez estimate this factor as 0.1
[7]
).
• is the fraction of stars in the Milky Way with planets.
• is the fraction of planets that are rocky ("metallic") rather than gaseous.
• is the fraction of habitable planets where microbial life arises. Ward and Brownlee believe this fraction is
unlikely to be small.
• is the fraction of planets where complex life evolves. For 80% of the time since microbial life first appeared
on the Earth, there was only bacterial life. Hence Ward and Brownlee argue that this fraction may be very small.
• is the fraction of the total lifespan of a planet during which complex life is present. Complex life cannot
endure indefinitely, because the energy put out by the sort of star that allows complex life to emerge gradually
rises, and the central star eventually becomes a red giant, engulfing all planets in the planetary habitable zone.
Also, given enough time, a catastrophic extinction of all complex life becomes ever more likely.
• is the fraction of habitable planets with a large moon. If the giant impact theory of the Moon's origin is
correct, this fraction is small.
• is the fraction of planetary systems with large Jovian planets. This fraction could be large.
• is the fraction of planets with a sufficiently low number of extinction events. Ward and Brownlee argue that
the low number of such events the Earth has experienced since the Cambrian explosion may be unusual, in which
case this fraction would be small.
The Rare Earth equation, unlike the Drake equation, does not factor the probability that complex life evolves into
intelligent life that discovers technology. (Keep in mind that Ward and Brownlee are not evolutionary biologists.)
Barrow and Tipler [30] review the consensus among such biologists that the evolutionary path from primitive
Cambrian chordates, e.g. Pikaia, to Homo sapiens was a highly improbable event. For example, the large brains of
humans have marked adaptive disadvantages, requiring as they do an expensive metabolism, a long gestation period,
and a childhood lasting more than 25% of the average total life span. Other improbable features of humans include:
• Being the only extant bipedal land (non-avian) vertebrate. Combined with an unusual eye–hand coordination, this
permits dextrous manipulations of the physical environment with the hands;
• A vocal apparatus far more expressive than that of any other mammal, enabling speech. Speech makes it possible
for humans to interact cooperatively, to share knowledge, and to acquire a culture;
• The capability of formulating abstractions to a degree permitting the invention of mathematics, and the discovery
of science and technology. Keep in mind how recently humans acquired anything like their current scientific and
technological sophistication.
Advocates
Books that advocate the Rare Earth hypothesis include:
• Stuart Ross Taylor [31] , a specialist on the solar system, firmly believes in the hypothesis, but its truth is not
central to his purpose, which is to write a short introductory book on the solar system and its formation. Taylor
concludes that the solar system is probably very unusual, because it resulted from so many chance factors and
events.
• Stephen Webb [2] , a physicist, mainly presents and rejects candidate solutions for the Fermi paradox. The Rare
Earth hypothesis emerges as one of the few solutions left standing by the end of the book.
• Simon Conway Morris [32] , a paleontologist, mainly argues that evolution is convergent. Morris devotes chapter
5 to the Rare Earth hypothesis, citing Rare Earth with approval. Yet while Morris agrees that the Earth could well
be the only planet in the Milky Way harboring complex life, he sees the evolution of complex life into intelligent
life as fairly probable, contra Ernst Mayr's views as reported in section 3.2 of the following reference.
• John D. Barrow and Frank J. Tipler (1986. 3.2, 8.7, 9), cosmologists, vigorously defend the hypothesis that
humans are likely to be the only intelligent life in the Milky Way, and perhaps the entire universe. But this
hypothesis is not central to their book, a very thorough study of the anthropic principle, and of how the laws of
physics are peculiarly suited to enable the emergence of complexity in nature.
• Ray Kurzweil, a computer pioneer and self-proclaimed Singularitarian, argues in The Singularity Is Near that the
coming Singularity requires that Earth be the first planet on which sentient, technology-using life evolved.
Although other Earth-like planets could exist, Earth must be the most evolutionarily advanced, because otherwise
we would have seen evidence that another culture had experienced the Singularity and expanded to harness the
full computational capacity of the physical universe.
Criticism
Criticisms of the Rare Earth Hypothesis take various forms.
Exoplanets are common

Over 400 exoplanets are known in late 2009 and more are continually discovered.[33] Alan Boss of the Carnegie
Institution of Science estimates there may be 100 billion terrestrial planets in our Milky Way Galaxy alone.[34] Boss
believes many could have simple lifeforms and there could be thousands of civilizations in our galaxy. He uses an
estimate that each sun-like star has on average one Earth-like planet.
Evolutionary biology
Central to the Rare Earth hypothesis is the following claim about evolutionary biology: while microbes of some sort
could well be common in the universe, complex life is unlikely to be. Yet to date, the only evolutionary biologist to
speak to the hypothesis at any length is Simon Conway Morris (2003). The hypothesis concludes, more or less, that
complex life is rare because it can evolve only on the surface of an Earth-like planet or on a suitable satellite of a
planet. Some biologists, such as Jack Cohen, believe this assumption too restrictive and unimaginative; they see it as
a form of circular reasoning (see Alternative biochemistry, a speculative biochemistry of alien life forms). Earth-like
planets may indeed be very rare, but non carbon-based complex life could possibly emerge in other environments.[35]
According to David Darling, the Rare Earth hypothesis is neither hypothesis nor prediction, but merely a description
of how life arose on Earth.[36] In his view Ward and Brownlee have done nothing more than select the factors that
best suit their case.
What matters is not whether there's anything unusual about the Earth; there's going to be something
idiosyncratic about every planet in space. What matters is whether any of Earth's circumstances are not
only unusual but also essential for complex life. So far we've seen nothing to suggest there is.
See also
• Astrochemistry
• Astrogeology
• Extrasolar planet
• Extraterrestrial life
• Evolving the Alien: The Science of Extraterrestrial Life
• Goldilocks planet
• History of Earth
• The mediocrity principle and cosmic pluralism are the antithesis of the Rare Earth hypothesis.
• Metaphysical naturalism
• Neocatastrophism
• Origin of life
• Panspermia
• Planetary habitability
• Precambrian
• Snowball earth
• Timetable of the Precambrian
References
• 'Hundreds of worlds' in Milky Way [37]
• [ Edit this reference [38]]
Barrow, John D.; Tipler, Frank J. (19 May 1988). The Anthropic Cosmological Principle [39]. foreword by John A.
Wheeler. Oxford: Oxford University Press. LC 87-28148 [40]. ISBN 9780192821478. Retrieved 31 December 2009.
• Cirkovic, Milan M., and Bradbury, Robert J., 2006, "Galactic Gradients, Postbiological Evolution, and the
Apparent Failure of SETI, [41]" New Astronomy, vol. 11, pp. 628–639.
• Comins, Neil F., 1993. What if the moon didn't exist? Voyages to Earths that might have been. HarperCollins.
• Simon Conway Morris, 2003. Life's Solution. Cambridge Univ. Press. See chpt. 5; many references.
• Cohen, Jack, and Ian Stewart, 2004 (2002). What Does a Martian Look Like: The Science of Extraterrestrial Life.
Ebury Press. ISBN 0-09-187927-2.
• Cramer, John G., 2000, "The 'Rare Earth' Hypothesis, [42]" Analog Science Fiction & Fact Magazine (September
2000).
• Guillermo Gonzalez, Brownlee, Donald, and Ward, Peter, 2001, "The Galactic Habitable Zone: Galactic
Chemical Evolution, [43]" Icarus 152: 185–200.
• James Kasting, Whitmire, D. P., and Reynolds, R. T., 1993, "Habitable zones around main sequence stars," Icarus
101: 108–28.
• Kirschvink, Joseph L., Robert L. Ripperdan, and David A. Evans, 1997, "Evidence for a Large-Scale
Reorganization of Early Cambrian Continental Masses by Inertial Interchange True Polar Wander, [44]" Science
277: 541–45.
• Knoll, Andrew H., 2003. Life on a Young Planet: The First Three Billion Years of Evolution on Earth. Princeton
Univ. Press.
• Dartnell, Lewis, 2007, Life in the Universe, a Beginner's Guide, One World, Oxford.
• Lineweaver, Charles H., Fenner, Yeshe, and Gibson, Brad K., 2004, "The Galactic Habitable Zone and the Age
Distribution of Complex Life in the Milky Way [45]," Science 303: 59–62.
• Lissauer, 1999, "How common are habitable planets?" Nature 402: C11-14.
• Prantzos, Nikos, 2006, "On the Galactic Habitable Zone [46]" in Bada, J. et al., eds., Strategies for Life Detection.
To appear in Space Science Reviews.
• Ross, Hugh, 1993, "Some of the parameters of the galaxy-sun-earth-moon system necessary for advanced life" in
The Creator and the Cosmos, 2nd ed. Colorado Springs CO: NavPress.
• Taylor, Stuart Ross, 1998. Destiny or Chance: Our Solar System and Its Place in the Cosmos. Cambridge Univ.
Press.
• Frank J. Tipler, 2003, "Intelligent Life in Cosmology," International Journal of Astrobiology 2: 141–48.
• Ward, Peter D., and Brownlee, Donald, 2000. Rare Earth: Why Complex Life is Uncommon in the Universe.
Copernicus Books (Springer Verlag). ISBN 0-387-98701-0.
• Webb, Stephen, 2002. Where is Everybody? (If the universe is teeming with aliens, Where is Everybody?: Fifty
solutions to the Fermi paradox and the problem of extraterrestrial life) Copernicus Books (Springer Verlag).
• Victor Stenger, 1999, "The Anthropic Coincidences: A Natural Explanation, [47]" The Skeptical Intelligencer 3(3).
External links
• Home page [48] of Rare Earth.
• Reviews of Rare Earth:
• Athena Andreadis [49], PhD in molecular biology.
• Kendrick Frazier [50], editor, Skeptical Inquirer.
• Tal Cohen [51], PhD student in computer science.
• "Galactic Habitable Zone, [52]" Astrobiology Magazine, May 18, 2001.
• Gregg Easterbrook, "Are We Alone? [53]" The Atlantic Monthly, August 1988. Article that anticipates REH in
some respects.
• Solstation.com: "Stars and Habitable Planets. [54]"
• Recer, Paul, "Radio astronomers measure sun's orbit around Milky Way, [55]" Associated Press, June 1, 1999.
• Breitbart.com, "Parallel universes exist, [56]" Sept. 23 2007.
References
[1] Brownlee and Ward (2000), pp. xxi–xxiii.
[2] Webb, Stephen, 2002. If the universe is teeming with aliens, where is everybody? Fifty solutions to the Fermi paradox and the problem of
extraterrestrial life. Copernicus Books (Springer Verlag)
[3] 1 Morphology of Our Galaxy's 'Twin' (http:/ / www. spitzer. caltech. edu/ Media/ releases/ ssc2004-12/ ssc2004-12a. shtml) Spitzer Space
Telescope, Jet Propulsion Laboratory, NASA.
[4] Brownlee, Donald. Ward, Peter D. "Rare Earth", pages 27 – 29. Copernicus. 2000.
[5] Lineweaver, Charles H., Fenner, Yeshe, and Gibson, Brad K., 2004, " The Galactic Habitable Zone and the Age Distribution of Complex Life
in the Milky Way (http:/ / astronomy. swin. edu. au/ GHZ/ GHZ_astroph. pdf)," Science 303: 59–62.
[6] Brownlee, Donald. Ward, Peter D. "Rare Earth", pages 32. Copernicus. 2000.
[7] Guillermo Gonzalez, Brownlee, Donald, and Ward, Peter, 2001, " The Galactic Habitable Zone: Galactic Chemical Evolution, (http:/ / adsabs.
harvard. edu/ cgi-bin/ nph-bib_query?bibcode=2001Icar. . 152. . 185G& db_key=AST& high=3c61d2959625890)" Icarus 152: 185–200.
[8] How often does the Sun pass through a spiral arm in the Milky Way? (http:/ / curious. astro. cornell. edu/ question. php?number=402), Karen
Masters, Curious About Astronomy
[9] Dartnell, Lewis, Life in the Universe, One World, Oxford, 2007, p. 75.
[10] Hart, M. "Habitable Zones Around Main Sequence Stars," Icarus, 37, 351 (1979).
[11] Reynolds, R. T., McKay, C. P., and Kasting, J. F. "Europa, Tidally Heated Oceans, and Habitable Zones Around Giant Planets," Advances
in Space Research, 7 (5), 125 (1987).
[12] James Kasting, Whitmire, D. P., and Reynolds, R. T., 1993, "Habitable zones around main sequence stars," Icarus 101: 108–28.
[13] Brownlee, Donald. Ward, Peter D. "Rare Earth", page 18. Copernicus. 2000.
[14] (http:/ / joy. chara. gsu. edu/ RECONS/ TOP100. posted. htm) The One Hundred Nearest Star Systems, Research Consortium on Nearby
Stars.
[15] Brownlee, Donald. Ward, Peter D. "Rare Earth", pages 15–33. Copernicus. 2000.
[16] Jupiter, entry in the Oxford English Dictionary, prepared by J. A. Simpson and E. S. C. Weiner, vol. 8, second edition, Oxford: Clarendon
Press, 1989. ISBN 0-19-861220-6 (vol. 8), ISBN 0-19-861186-2 (set.)
[17] Horner, J.; Jones, B.W. (2008). "Jupiter – friend or foe? I: the asteroids" (http:/ / xxx. lanl. gov/ ftp/ arxiv/ papers/ 0806/ 0806. 2795. pdf).
International Journal of Astrobiology 7 (3&4): 251–261. doi:10.1017/S1473550408004187. . Retrieved 2009-05-15.
[18] Hinse, T.C.. "Chaos and Planet-Particle Dynamics within the Habitable Zone of Extrasolar Planetary Systems (A qualitative numerical
stability study)" (http:/ / www. astro. ku. dk/ ~tobiash/ posters/ Tobias_final_new. pdf) (PDF). Niels Bohr Institute. . Retrieved 2007-10-31.
"Main simulation results observed: [1] The presence of high-order mean-motion resonances for large values of giant planet eccentricity [2]
Chaos dominated dynamics within the habitable zone(s) at large values of giant planet mass."
[19] "Once you realize that most of the known extrasolar planets have highly eccentric orbits (like the planets in Upsilon Andromedae), you
begin to wonder if there might be something special about our solar system" (UCBerkeleyNews quoting Extra solar planetary researcher Eric
Ford.) Sanders, Robert (13 April 2005). "Wayward planet knocks extrasolar planets for a loop" (http:/ / www. berkeley. edu/ news/ media/
releases/ 2005/ 04/ 13_planet. shtml). . Retrieved 2007-10-31.
[20] Lissauer 1999, as summarized by Conway Morris 2003: 92; also see Comins 1993
[21] Planet-hunters set for big bounty (http:/ / news. bbc. co. uk/ 1/ hi/ sci/ tech/ 7249884. stm), BBC
[22] Taylor 1998
[23] Dartnell, pp. 69–70,
[24] A formal description of the hypothesis is given in: Lathe, Richard (March 2004). "Fast tidal cycling and the origin of life". Icarus 168 (1):
18–22. doi:10.1016/j.icarus.2003.10.018. "tidal cycling, resembling the polymerase chain reaction (PCR) mechanism, could only replicate and
amplify DNA-like polymers. This mechanism suggests constraints on the evolution of extra-terrestrial life.". It is taught less formally here:
Schombert, James. "Origin of Life" (http:/ / abyss. uoregon. edu/ ~js/ ast121/ lectures/ lec25. html). University of Oregon. . Retrieved
2007-10-31. "with the vastness of the Earth's oceans it is statistically very improbable that these early proteins would ever link up. The
solution is that the huge tides from the Moon produced inland tidal pools, which would fill and evaporate on a regular basis to produce high
concentrations of amino acids".
[25] "New Map Provides More Evidence Mars Once Like Earth" (http:/ / www. nasa. gov/ centers/ goddard/ news/ topstory/ 2005/ mgs_plates.
html). 10 December 2005. .
[26] Belbruno, E.; J. Richard Gott III (2005). "Where Did The Moon Come From?". The Astronomical Journal 129 (3): 1724–1745.
doi:10.1086/427539. arXiv:astro-ph/0405372.
[27] Taylor, Stuart Ross, 1998. Destiny or Chance: Our Solar System and Its Place in the Cosmos. Cambridge Univ. Press
[28] Cramer (2000)
[29] Brownlee, Donald. Ward, Peter D. "Rare Earth", pages 271–275. Copernicus. 2000.
[30] [ Edit this reference (http:/ / en. wikipedia. org/ w/ index. php?title=Template:BarrowTipler1986& action=edit)]
Barrow, John D.; Tipler, Frank J. (19 May 1988). The Anthropic Cosmological Principle (http:/ / books. google.
com/ books?id=uSykSbXklWEC& printsec=frontcover). foreword by John A. Wheeler. Oxford: Oxford University
Press. LC 87-28148 (http:/ / lccn. loc. gov/ 87028148). ISBN 9780192821478. . Retrieved 31 December 2009.
Section 3.2
[31] Taylor, Stuart Ross, 1998. Destiny or Chance: Our Solar System and Its Place in the Cosmos. Cambridge Univ. Press.
[32] Simon Conway Morris, 2003. Life's Solution. Cambridge Univ. Press. See chpt. 5; many references.
[33] Schneider, Jean. "Interactive Extra-solar Planets Catalog" (http:/ / exoplanet. eu/ catalog. php). The Extrasolar Planets Encyclopedia. .
[34] Galaxy may be full of 'Earths,' alien life (http:/ / www. cnn. com/ 2009/ TECH/ space/ 02/ 25/ galaxy. planets. kepler/ )
[35] For a detailed critique of the Rare Earth hypothesis along these lines, see Cohen and Ian Stewart (2002).
[36] Darling, David (2001). Life Everywhere: The Maverick Science of Astrobiology. Basic Books/Perseus.
[37] http:/ / news. bbc. co. uk/ 1/ hi/ sci/ tech/ 7249884. stm
[38] http:/ / en. wikipedia. org/ w/ index. php?title=Template:BarrowTipler1986& action=edit
[39] http:/ / books. google. com/ books?id=uSykSbXklWEC& printsec=frontcover
[40] http:/ / lccn. loc. gov/ 87028148
[41] http:/ / www. anthropic-principle. com/ preprints/ milan-seti. pdf
[42] http:/ / www. npl. washington. edu/ av/ altvw102. html
[43] http:/ / adsabs. harvard. edu/ cgi-bin/ nph-bib_query?bibcode=2001Icar. . 152. . 185G& db_key=AST& high=3c61d2959625890
[44] http:/ / www. sciencemag. org/ cgi/ content/ abstract/ 277/ 5325/ 541
[45] http:/ / astronomy. swin. edu. au/ GHZ/ GHZ_astroph. pdf
[46] http:/ / arxiv. org/ abs/ astro-ph/ 0612316
[47] http:/ / www. colorado. edu/ philosophy/ vstenger/ Cosmo/ anthro_skintel. html
[48] http:/ / www. astro. washington. edu/ rareearth/
[49] http:/ / www. setileague. org/ reviews/ rarearth. htm
[50] http:/ / www. findarticles. com/ p/ articles/ mi_m2843/ is_6_25/ ai_79794362
[51] http:/ / tal. forum2. org/ rare
[52] http:/ / www. astrobio. net/ news/ modules. php?op=modload& name=News& file=article& sid=139
[53] http:/ / www. theatlantic. com/ issues/ 88aug/ easterbr. htm
[54] http:/ / www. solstation. com/ habitable. htm
[55] http:/ / www. chron. com/ content/ interactive/ space/ astronomy/ news/ 1999/ ds/ 990602. html
[56] http:/ / www. breitbart. com/ article. php?id=paUniverse_sun14_parallel_universes& show_article=1& cat=0
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Image Sources, Licenses and Contributors 67
Image Sources, Licenses and Contributors

Image:BlueMarble-2001-2002.jpg Source: http://en.wikipedia.org/w/index.php?title=File:BlueMarble-2001-2002.jpg License: Public Domain Contributors: User:Apoc2400
Image:The Earth seen from Apollo 17.jpg Source: http://en.wikipedia.org/w/index.php?title=File:The_Earth_seen_from_Apollo_17.jpg License: Public Domain Contributors: NASA. Photo
taken by either Harrison Schmitt or Ron Evans (of the Apollo 17 crew).
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Origin of Life

Origin of water on Earth

Origins Water covers about 70% of the Earth's surface

Some of the most likely contributory factors

Water in the development of the Earth

Suitable star systems

A stable habitable zone

A "stable" HZ denotes two factors.

Low stellar variation

Orbit and rotation

Microenvironments and extremophiles

The discovery of life in extreme conditions has complicated definitions

Alternative star systems

Red dwarf systems

Astronomers for many years ruled out red

little light (from 3% of that produced by the

The galactic neighborhood

Life's impact on habitability

be within the HZ—this remains to be proven.

Pasteur and Darwin

In a letter to Joseph Dalton Hooker on February 1, 1871,[14] Charles Darwin

"Primordial soup" theory

Origin of organic molecules

"Soup" theory today: Miller's experiment and subsequent work

Regarding the reducing atmosphere

Regarding monomer formation

Regarding monomer accumulation

Regarding further transformation

The deep sea vent theory

Another possible answer to this polymerization conundrum was

In contrast to the classical Miller experiments, which depend on

Radioactive beach hypothesis

Thermodynamic Origin of Life: Ultraviolet and Temperature-Assisted Replication (UVTAR) Model

Models to explain homochirality

bias, whereas sugars show a predominantly right-handed bias.[56]

Self-organization and replication

From organic molecules to protocells

"Genes first" models: the RNA world

"Metabolism first" models

Possible role of bubbles

Gold's "Deep-hot biosphere" model

"Primitive" extraterrestrial life

Extraterrestrial amino acids

PAH world hypothesis

The Miller–Urey experiment[1] (or

In 2008,[7] a re-analysis of Miller's The experiment

archived solutions from the original

Earth's early atmosphere

Recent related studies

Entropy and life

The solar system from a thermodynamic systems perspective.

that system is entirely compatible with the second law.[5]

Gibbs free energy and biological evolution

Entropy and the origin of life

Group: Group I (dsDNA)

(unranked): Nucleocytoplasmic large DNA viruses

Species: Acanthamoeba polyphaga mimivirus

the host cell.

Implications for defining "life"

References in popular culture

[22] http:/ / www. expasy. org/ viralzone/ all_by_species/ 17. html

Rare Earth hypothesis