O

See also
O2 ▶ Oceanic Crust
▶ Ophiolite
▶ Dioxygen ▶ Plate Tectonics
▶ Subduction

OB Association
Definition
Oberon
An OB association is a loose grouping of several thousand
Definition
stars, a small fraction of which are of spectral type O and B.
Oberon, discovered by Herschel in 1787, is the outermost
These latter are the most massive members of the group and
of the five big satellites of ▶ Uranus. Its distance to Uranus
also the most luminous, allowing identification of OB asso-
is 583,500 km (or 23 Uranian radii), its diameter is
ciations out to great distances. In external spiral galaxies,
1,520 km, and its density is 1.63 g/cm3. Oberon has been
they trace the spiral arms. OB associations range in size, with
observed by the ▶ Voyager 2 spacecraft at the time of its
diameters from tens of parsecs to about 100 parsecs. It is
▶ Uranus flyby in January 1986. Its surface is heavily
thought that all the stars formed together in a tighter con-
cratered; the largest crater, Hamlet, has a diameter of
figuration, and observations of stellar velocities reveal that
206 km. It is considered as being differentiated into a sili-
the groups are in a state of expansion. The oldest recogniz-
cate core surrounded by an icy mantle. The low albedo
able OB associations have ages of order 10 million years.
(0.14) of the surface suggests that it consists of a mixture
of ice and organic matter, possibly irradiated by high-
See also energy particles coming from Uranus’ magnetosphere.
▶ Elephant Trunks
▶ Pre-Main-Sequence Star
▶ Stellar Cluster
See also
▶ Giant Planets
▶ T Association
▶ Uranus
▶ Voyager

Obduction
Definition Obliquity and Obliquity
Obduction is the thrusting of segments of ▶ oceanic crust Variations
and mantle onto continental crust at convergent plate
margins. A slice of obducted oceanic crust is called an FRANÇOIS FORGET
▶ ophiolite. Commonly, blueschist-facies metamorphic Institut Pierre Simon Laplace, Laboratoire de
rocks are exposed in front of the obducted ophiolite. Dur- Météorologie Dynamique, UMR 8539, Université Paris 6,
ing periods of enhanced ridge push, while most of the Paris Cedex 05, France
oceanic plate is subducted, slices of more buoyant material
are thrust onto small continental fragments or onto the
margins of continents, particularly during the closure of Synonyms
ocean basins preceding continent–continent collision. Axial tilt

Muriel Gargaud (ed.), Encyclopedia of Astrobiology, DOI 10.1007/978-3-642-11274-4,

# Springer-Verlag Berlin Heidelberg 2011
1160 O Occultation

Definition about 15
and 35
. Between 5 and 20 million years ago,
In astronomy, the obliquity is the angle between an it oscillated between 25
and 45
. Before that, it is
object’s (e.g., planet’s) axis of ▶ rotation and a line per- impossible to reconstruct the details of the obliquity
pendicular to its ▶ orbit plane. The obliquity controls the variations, because of the chaotic nature of the problem
variation of insolation with latitude and time, and thus (Laskar et al. 2004).
influences the climate. On any planet, the obliquity controls the distribution
of solar illumination with latitude, and the seasonal cycle.
Overview Increasing the obliquity enhances the seasonal variations
In the Solar System, the present-day obliquities of the (warmer summer, colder winter). The average annual
▶ planets are diverse Table 1. insolation slightly decreases with obliquity equatorward
Planetary obliquities vary with time because of the of 45
latitude, and strongly increases with obliquity pole-
long-term gravitational perturbation by other planets ward of 45
latitude. At the poles, it is proportional to the
(Laskar and Robutel, 1993). This effect is small on the sine of the obliquity.
obliquity of the outer planets (▶ Jupiter, ▶ Saturn, ▶ Ura-
nus and ▶ Neptune), which can be considered as primor- See also
dial: They have roughly kept the value they had at the end ▶ Earth
of the formation of the ▶ Solar System. In contrast, the ▶ Jupiter
very low obliquity of ▶ Mercury results from tidal inter- ▶ Mercury
actions with the Sun. This is also the case for ▶ Venus, ▶ Moon, The
although the effect was different because of its thick atmo- ▶ Neptune
sphere (the Sun heats the atmosphere at the subsolar point, ▶ Orbit
inducing a redistribution of the mass of the atmosphere, ▶ Planet
which also induces a torque). For the ▶ Earth, the obliquity ▶ Rotation Planet
presents only small variations of about 1.3
around the ▶ Saturn
mean value of 23.3
, with a dominating period of 42,000 ▶ Solar System, Inner
years. In the absence of the ▶ Moon, theoretical calcula- ▶ Uranus
tions suggest that the situation would be very different, as ▶ Venus
multiple resonances then occur between the precession of
the axis and the precession of the orbital plane. Earth’s References and Further Reading
obliquity would then have varied widely between nearly Laskar J, Robutel P (1993) The chaotic obliquity of the planets. Nature
0
and 85
. In fact, this is the case for Mars, which 361:608–612
may have varied between 0
and up to more than 60
. Laskar J, Correia ACM, Gastineau M, Joutel F, Levrard B, Robutel P
(2004) Long term evolution and chaotic diffusion of the insolation
In the past 5 million years, the Martian obliquity varied
quantities of Mars. Icarus 170:343–364
(with a pseudo-period near 120,000 Earth years) between

Obliquity and Obliquity Variations. Table 1 Obliquity of the Occultation

eight planets of the Solar System and Pluto

Planet Obliquity Definition

In astronomical usage, an occultation occurs when an
Mercury 0.01

object of larger angular size passes in front of an object
Venus 177.4
of smaller angular size. A frequent example is the passage
Earth 23.45
of the Moon between an observer on Earth and a star,
Mars 25.19
producing an occultation of the star. The rarer occulta-
Jupiter 3.1
tions of stars by planets, satellites, or asteroids provide
Saturn 26.7
opportunities to probe the properties (atmosphere, size)
Uranus 97.8
of the occulting bodies. Such observations led to the
discovery of the rings around the planets ▶ Uranus and
Neptune 28.3

▶ Neptune. When the Earth passes through the equatorial
Pluto 122.5

plane of ▶ Jupiter or ▶ Saturn (which correspond to the
 Ocean Planet O 1161

orbital planes of the major satellites of these planets), Definition

mutual occultations among these satellites can be observed. Ocean planets are hypothetical objects, formed beyond the
Occultation events have also been observed from unmanned snow line of a planetary system (distance in the proto-
space probes in the Jovian and Saturnian systems. The case planetary nebula at which the temperature is cool enough
when the foreground object has a smaller angular size than to allow the hydrogen compounds such as H2O, NH3, CH4
the more distant object is called a ▶ transit. to condense, essentially when the gas temperature reaches
the sublimation temperature of water, 170 K in the vac-
See also uum; see Hayashi 1981). Their mass between about 5 and
▶ Eclipse 10 terrestrial masses would be made of volatiles and of
▶ Jupiter silicate rocks and iron. In the prototype model, the pro-
▶ Neptune portion was 50%–50%, which corresponds to the maxi-
▶ Saturn mum ratio between volatiles and rocks found in the Solar
▶ Transit System objects (except for some water-dominated satel-
▶ Uranus lites of Saturn), but 1–10% of volatiles may be sufficient to
form an ocean planet. Following orbital migration, the
snow line is crossed by the planet and, after orbital stabi-
lization within the ▶ habitable zone of the planetary sys-
Ocean (on Early Venus) tem, part of the volatiles become liquid, forming a deep
ocean at the surface of the planet.
Definition
Liquid water oceans may have existed on early ▶ Venus, History
when the solar luminosity was weaker than today. The The concept of ocean planet was first discussed in 2003
ocean would have disappeared no later than about 1 bil- and led to two publications (Kuchner 2003; Léger et al.
lion years ago, the epoch of global Venus resurfacing. The 2004). Ocean planets are now considered as particular
existence of an ocean remains speculative, since no direct cases in the larger framework of rocky planets’ composi-
observational evidence is available. The observation of tion and structure (Valencia et al. 2007), as opposed to the
a 100-fold enrichment in ▶ deuterium in the Venus atmo- gas giant planets.
sphere has been frequently cited as indirect evidence of the
presence of significant amounts of ▶ water on early Venus, Overview O
though. The concept of ocean planet is the result of a thought exper-
iment. In the classical theory of planetary formation (e.g.,
See also Wuchterl et al. 2000), accretion of planets beyond the snow
▶ Deuterium line is rapid because volatiles in solid form provide more
▶ Venus quickly sufficient mass of material to build the core of the
▶ Water planet. Some of these formed planetary cores are massive
enough to accrete gas and form giant planets. Some are
not and lead to small frozen objects such as Pluto or icy
satellites of giant planets in our Solar System. The thresh-
Ocean Planet old is estimated to be around 6–10 Earth masses; but this
value depends on the disk density and the orbital distance.
MARC OLLIVIER In some case, atmospheric effects such as trapping of the
Institut d’Astrophysique Spatiale, CNRS, Université de gas by a cold trap should also be considered.
Paris-Sud, Orsay, France Let us consider one of these several Earth-mass frozen
planet. Migration can bring this planet inward, trespassing
the snow line (Goldreich and Tremaine 1980), and stabi-
Synonyms lizing the planetary orbit, possibly within its habitable zone.
Water planet; Water world This new position leads part of the volatiles to melt.
According to the planetary final semi-major axis, the authors
Keywords of this theory proposed that part of the volatiles can stay in
Accretion, exoplanet, migration, planetary formation, the liquid state at the surface of the planet, creating thus
snow line a huge and deep planetary ocean. However, several questions
1162 O Ocean Planet
have been raised concerning the internal structure of such
planets, the real existence of a huge ocean and its potential
characteristics, and the composition of the atmosphere
allowing its remote identification and study as an exobiology
object. At present, only Léger et al. (2004) have published
a specific study on these objects. As mentioned before,
other studies have been done in a larger context of terres-
trial-type planet structure (that thus includes ocean
planets.) In this entry we will consider Léger et al.’s work.
Several supplementary assumptions as limits of their
model have been done (Léger et al. 2004):
● The mass range of planets is limited to 1–8 Earth
masses. As biggest objects are easier to detect and
characterize, the study focuses on 6–8 Earth mass-
objects.
● Planets are supposed to form beyond the snowline and
migrate inward to about 1 AU (habitable zone of a
G star) in about 1 Myr.
● Planets are mostly made of 50% of refractory materials
(Fe, silicates) and 50% of ices according to the com-
position of protoplanetary nebula (90% of H2O, 5% of
NH3, 5% of CO2). Most of the ice is thus made of
water.
Internal Structure
Modeling of a 6-Earth mass-object has been done starting
from a model developed to describe the Earth’s interior,
taking into account the cooling of the planet, and the
e ∼ 100 km
R/R =
2.00
1.24
0.69
6 M⊕ Ocean-Planet 6 M⊕ Rocky Planet
(3 M⊕ metals and silicates (6 M⊕ metals and silicates)
+ 3 M⊕ H2O)
Ocean Planet. Figure 1 Calculated internal structure of a 6-Earth mass-ocean-planet (left). The core (Fe), mantle (silicates),
and ice + ocean masses are 1, 2, and 3 Earth masses, respectively. Comparison is made with a 6-Earth mass-fully rocky planet
(centre) and the Earth (right) (From Léger et al. 2004)
composition and structure of ice at such extreme pressure
and temperature values. The structure of such an object is
described in Fig. 1.
The ocean, with a thickness of about 100 km, lies on
a huge water ice layer (several thousands of kilometers).
Because of the very high pressure, ice is differentiated.
Because of its intrinsic weight compared to water ice,
CO2 is mainly in the form of ice, at the interface between
the mantle and the water ice layer. This helps prevent CO2
from degassing into the atmosphere, creating to a runaway
greenhouse effect.
Ocean Depth
The question of the depth of the ocean was studied con-
sidering a pure water ocean, whose role is to transfer the
internal heat of the planet to its surface. Assuming the
worst case where the temperature gradient is adiabatic
(limit to convection), a surface temperature of the ocean
of 7
C is obtained. The depth of the ocean is estimated at
72 km. This value is higher if the surface temperature of
the ocean is higher and vice versa. Non-adiabatic depen-
dence of the temperature with depth leads to a smaller
ocean depth.
Atmosphere Composition and Spectrum
Our understanding of the origin of atmospheres (includ-
ing the terrestrial one) is too poor to make accurate pre-
dictions of what an ocean planet atmosphere would
exactly contain. However, assuming the composition of
1.63
e ∼ 20 km
1.00
0.85
0.55
Earth
(1 M⊕ metals and silicates)

cometary material, it is not irrelevant to assume that
volatiles are made of H2O (mainly), NH3, and CO2. In
the atmosphere, NH3 reacts with EUV radiations and
dissociates leading to H2 and N2. Because of high exo-
sphere temperature, H2 escapes and N2 is mixed with the
remaining atmosphere as a buffer gas. The amount of CO2
in the atmosphere is strongly limited compared to the
planetary CO2, because most of the CO2 is in the form
of ice under the water ice layer. Models of ocean planet
atmospheres including photochemical evolution include
thus the presence of H2O, CO2, and N2 that should be
detectable in the atmosphere.
Exobiological Interest of such Planets
Because of the massive presence of liquid water at their
surface, ocean planets are good candidates for the detec-
tion of life, assuming the classical criteria for the potential
emergence of life (definition of the habitable zone).
A search for life on these planets requires first an unam-
biguous identification of their nature (see next section),
coming from the determination of their mean density, and
the confirmation of their habitability thanks to a spectral
analysis of their atmosphere. However, because of the lack
of O2 sinks (such as the Earth’s silicate surface), it appears
possible to maintain a quantity of abiotic O2 coming from
photodissociation of CO2 or H2O in the atmosphere. The
CO2, O2, H2O criteria cannot thus be used. However, in
that case, the formation of a large ozone (O3) layer
is limited by perpetual reaction of O3 in the upper
atmosphere with radicals coming from H2O and CO2
dissociation. The effective habitability of ocean planets
could thus be confirmed by searching for a well-developed
ozone layer, which implies the occurrence of a huge
amount of biologically produced O2 in the atmosphere.
Key Research Findings
No observational confirmation of the existence of ocean
planets has yet been done. To confirm the existence of
these objects, one must determine, with sufficient accu-
racy to exclude all the other models, the density of the
object, which depends on its internal composition (vola-
tiles, iron, and silicates). In addition, mass-radius degen-
eracies should be taken into account carefully. A planet
with a dense core and a hydrogen-rich atmosphere cannot
certainly be distinguished from a H2O-rich planet, but
populations of such objects might be separated by the
fact that even low-mass hydrogen-rich envelopes produce
very large radii that cannot be attributed to ocean planets.
The amount of H2 that can mimic an ocean planet may
not be stable, assuming atmospheric escape.
Ocean Planet O 1163
To determine the density of the planet, one must
measure its radius and evaluate its mass. The first param-
eter can be obtained by the transit method. Observation
from space by dedicated satellites (▶ CoRoT, ▶ Kepler,
HST, etc) allows the determination of a planetary diameter
at the level of a few percent, assuming simultaneously the
knowledge of the stellar diameter with an accuracy of a few
percent. In principle, the mass of the planet can be deter-
mined by ▶ radial velocity measurements (RV) based on
high spectral resolution spectroscopy. However, the accu-
racy reached by the best RV instruments does not yet allow
measuring the mass of such small objects at a level better
than several Earth masses, preventing from determination
of the accurate density of the object. In that context, it is
thus very difficult to identify within a large range of
telluric planet models which one corresponds to the
observed planet, preventing the observer from identifying
a real ocean planet (Selsis et al. 2007). Future generations
of RV instrument should be more adapted to this quest.
The knowledge of stellar diameter may also be improved
by interferometric observations.
Future Directions
The concept of ocean planet is now considered as
a particular case of the bigger family of rocky planets
(mass between 2 and 10 terrestrial masses), whose
structure and evolution is driven by several parameters
such as the initial composition, the formation location,
O
and the evolution of orbital parameters. The observational
study of ocean planets is thus considered within the
framework of detection and characterization of telluric
planets.
See also
▶ CoRoT Satellite
▶ Exoplanet, Detection and Characterization
▶ Habitable Planet (Characterization)
▶ Habitable Zone
▶ Kepler Mission
▶ Radial Velocity
References and Further Reading
Goldreich P, Tremaine S (1980) Disk-satellite interactions. Astrophys J
241:425–441
Hayashi C (1981) Structure of the solar nebula, growth and decay of
magnetic fields and effects of magnetic and turbulent viscosities on
the nebula. Prog Theor Phys Suppl 70:35–53
Kuchner MJ (2003) Volatile-rich earth mass planets in the habitable zone.
Astrophys J 596:L105–L108
Léger A, Selsis F, Sotin C, Guillot T, Despois D, Mawet D, Ollivier M,
Labèque A, Valette C, Brachet F, Chazelas B, Lammer H (2004) A new
family of planets? “Ocean Planets”. Icarus 169:499–504
1164 O Ocean Salinity

Selsis F, Chazelas B, Bordé P, Ollivier M, Brachet F, Decaudin M, Bouchy F, Overview

Ehrenreich D, Griessmeier J-M, Lammer H, Sotin C, Grasset O,
Moutou C, Barge P, Deleuil M, Mawet D, Despois D, Kasting JF,
Léger A (2007) Could we identify hot ocean-planets with CoRoT,
Kepler and Doppler velocimetry. Icarus 191:453–468
Valencia D, Sasselov DD, O’Connell RJ (2007) Detailed models of
super-earths: how well can we infer bulk properties? Astrophys J
665:1413–1420
Wuchtel G, Guillot T, Lissauer JJ (2000) Giant planet formation. In:
Mannings V, Boss AP, Russell SS (eds) Protostars and Planets IV.
University of Arizona Press, Tucson, p 1081
Ocean Salinity
▶ Ocean, Chemical Evolution of
Ocean, Chemical Evolution of
DANIELE L. PINTI
GEOTOP & Département des Sciences de la Terre et de
l’Atmosphère, Université du Québec à Montréal,
Montréal, QC, Canada
Synonyms
Ocean salinity
Keywords
Major ions, ocean chemistry, oxygen levels, pH, salinity
Definition
The chemical composition of the ocean is the relative
compositions of major ions (Naþ, Mg2þ, Ca2þ, Kþ, Sr2þ,
Cl, SO2   2
4 , HCO3 , Br , CO3 , B(OH)3, B(OH)4 , F ) of
 
seawater. The major ions content is considered relatively
constant and defined as salinity, which is a measure of the
total dissolved salts in seawater (Table 1). The chemical
evolution of the ocean is the evolution of its chemistry
(major ions, pH, and oxygen content) and the processes
that controlled the composition of this terrestrial reservoir.
Though the chemical composition of the terrestrial oceans
is constant, a number of processes can cause the oceanic
waters to be nonconservative. These processes include
precipitation, dissolution, water–rock interactions, freez-
ing, and oxidation processes (Millero 2003). The terres-
trial oceans have certainly undergone an evolution of their
chemistry during the last 4.5 Ga (Holland 1984, 2003)
driven by the competition between two sources of salinity:
weathering of continental masses versus water cycling
through mid-ocean ridges (▶ MOR). Weathering of con-
tinental rocks and river waters mainly controls salinity of
present-day oceans. Seawater cycling at MOR was possibly
the dominant source of ions in the ▶ Hadean and
▶ Archean oceans. This would be a direct consequence
of a much vigorous mantle convection favoring higher
oceanic crust production at MOR (Bickle 1986) and the
limited growth of continents (McCulloch and Bennett
1994). Several authors indicated the beginning of the
Proterozoic era as the turning point in the dominance of
salinity sources (hydrothermal cycling versus continental
weathering; Veizer et al. 1989; de Ronde et al. 1997; Pinti
2005). The long geological history of seawater started in
the Hadean, when liquid water became stable at the sur-
face of the planet, 4.4–4.3 Ga. These ages correspond to
those of the ▶ zircons of ▶ Jack Hills (Wilde et al. 2001).
At present, little can be said about the composition of the
Hadean ocean (4.5–3.8 Ga) and mostly is pure specula-
tion. The chemistry of seawater in the early Hadean was
possibly controlled by high-temperature water–rock inter-
actions between the condensing CO2–H2O runaway
greenhouse atmosphere and the primitive basaltic crust.
When condensing water started to penetrate deeper in the
oceanic basaltic proto-crust, dissolved chlorine (degassed
form the mantle as HCl; Holland 1984) reacted with
Na-bearing minerals of basalt to produce halite (NaCl).
Sleep et al. (2001) calculated that a global basaltic layer
only 500-m thick could account for the whole ocean NaCl
budget. Initial salinity of the ocean is unknown and values
of 1.2–2 present value have been proposed only for the
Archean ocean (Holland 1984; Knauth 2005). These could
be considered as minimum values for the Hadean. The
presence of large amount of CO2 in the Hadean

Ocean, Chemical Evolution of. Table 1 Standard mean seawater chemical composition (Salinity = 35 g/L) reported as mol kg
H2O1 (DOE 1994)

Na+ Mg2+ Ca2+ K+ Sr2+ Cl SO42 HCO3 Br CO32 B(OH)3 B(OH)4 F
0.48616 0.05475 0.01065 0.01058 0.00009 0.56567 0.02927 0.00183 0.00087 0.00027 0.00033 0.00010 0.00007

atmosphere and oceans and large quantity of Na in the
basaltic proto-crust could have promoted their alkalinity
by production of Na2CO3, as observed in ▶ soda lakes
(Kempe and Degens 1985). However, thermodynamic cal-
culations indicated that the whole terrestrial available CO2
budget (atmosphere–hydrosphere–mantle) is several
orders of magnitude lower than the one needed to make
a global soda ocean (Sleep et al. 2001).
Seawater geological record started in the Archean,
though fragmentary and often hampered by large hydro-
thermal alteration and metamorphism. The 3.8 Ga ▶ Isua
metasedimentary sequences of West Greenland contain
chemical sediments deposited in an oceanic environment
without a significant sialic detrital component (Holland
2003), i.e., without continental inputs of ionic species.
Compelling evidence for the existence of an ocean during
deposition of Isua rocks are clastics that have been
interpreted as marine turbidites and iron formations
(IFs). These iron-rich layers are magnetite–quartz IFs
and the molar ratio of Fe2O3/FeO is less than 1.0, which
suggests that magnetite was the dominant ▶ iron oxide,
and that hematite was absent or very minor in these IFs
(Holland 2003). This in turn suggests that magnetite is not
a replacement of oxyhydroxides precursors but a primary
precipitating phase, thus that the oxygen level in the
oceans was insignificantly low. The oxidation of Fe2þ to
Fe3þ was possibly produced by UV-induced reactions
between iron and H2O (Cairns-Smith 1978). The
chemistry of Isua seawater is unknown. Appel et al.
(2001) observed ▶ fluid inclusions in quartz globules
from pillow breccia at Isua. The chemistry of the highly
saline aqueous fluids (about 25 wt.% NaCl equivalent)
bears a strong resemblance to present-day seafloor hydro-
thermal fluids that likely provoked the alteration of the
pillow breccia and the precipitation of quartz. Occur-
rences of fluid inclusions in Archean metasedimentary
rocks see Fluid Inclusions have been signaled in several
terrains of ages spanning from 3.5 to 2.7 Ga (De Ronde
et al. 1997; Channer et al. 1997; Foriel et al. 2004;
Weiershauser and Spooner 2005). They have been
interpreted as a pristine record of the chemical composi-
tion of Archan seawater. The chemistry of these fluid
inclusions always pointed out to the mixing between at
least two fluids, a saline fluid of Na–Ca–Cl composition
(seawater) and a hydrothermal effluent, often richer in
Ba and Fe (de Ronde et al. 1997; Foriel et al. 2004). The
Na:Cl ratio is the same as present-day seawater but NaCl
concentration ranges from 4% to 25% equivalent (against
3.5% equivalent in modern seawater). The salt enrichment
and the chemistry (Na–Ca–Cl) points out to an evaporate
seawater residue (Martin et al. 2006). Among the notable
Ocean, Chemical Evolution of O 1165
differences with modern seawater, de Ronde et al. (1997)
and Channer et al. (1997) noticed that the Cl:Br and Cl:I
ratios are higher than in modern seawater and much closer
to the bulk Earth values. This could indicate that Archean
seawater chemistry was buffered by the mantle through
emission of dissolved chemical species and volatiles from
mid-oceanic vents. An alternative hypothesis is the
absence of planktonic organisms that presently use bro-
mine and iodine as metabolites, scavenging them from
the ocean.
Holland (1984, 2003) used a different approach for
constraining the Archean seawater composition, looking
at the mineralogy of the deposited sediments. Calcite
(CaCO3) and dolomite (CaMg(CO3)2) were the dominant
carbonate minerals. ▶ Siderite (FeCO3) was only a
common constituent of ▶ BIFs. These observations
imply that the Archean ocean was supersaturated with
respect to CaCO3 and CaMg(CO3)2. For the theoretical
range of values of atmospheric pCO2 in the Archean, the
pH of seawater was probably between 6 and 7 (Holland
2003; Pinti 2005). The scarcity of siderite in BIFs is also an
indicator of a low Fe2þ/Ca2þ ratio in the ocean, i.e., close
to the ratio of the solubility product of siderite and calcite
(4  10–3; Holland 2003). Archean ocean should have been
mainly anoxic as suggested by the mass-independent frac-
tionation (▶ MIF) of the ▶ sulfur isotopes in pre-2.47 Ga
sulfides and sulfates (Farquhar et al. 2000). In the absence
of O2, solar UV interacts with SO2 and generates MIF of
the sulfur isotopes in the reaction products. In the absence O
of atmospheric O2, sulfide minerals would not have been
oxidized during weathering reducing the input of SO2 4 in
the ocean. Another consequence of the low pO2 in the
Archean was the low amount of trace elements such as U,
Mo, and Re that are extremely mobiles in an oxygenated
environment.
The Proterozoic ocean was certainly characterized by
an increasing O2 concentration, although it reached
appreciable amounts only after 1.8 Ga. The presence of
O2 certainly favored the scavenging of iron from the ocean
as oxyhydroxides and the deposition of ▶ BIFs (Holland
1984) and the increasing amount of sulfate. The cessation
of BIFs deposition after 1.8 Ga was probably due to
a decrease in the flux of reductants Fe2þ, H2, and H2S to
the oceans, which in turn promoted a net increase in the
dissolved O2 content in the deeper ocean (Holland 2006).
The end of the Proterozoic was a quite turbulent period
of biological and climatic global changes in the ocean
driven by very large glaciations that possibly developed
in ▶ Snowball Earths.
Phanerozoic is the period where the largest (but
discontinuous) geological record of the chemical
1166 O Ocean, Chemical Evolution of
evolution of the ocean is available. First studies pointed
out the near-constancy of the seawater chemical compo-
sition (e.g., Holser 1963; Holland 1972), which turned out
to be off the mark. Indeed, Phanerozoic is characterized by
large excursions in the concentration of several major
ions, particularly Ca2þ, Mg2þ, and SO2 4 , which contrast
with a nearly constancy of Kþ (Spencer and Hardie 1990;
Hardie 1996; Horita et al. 2002). These variations (2–5
the present seawater) have been differently interpreted, yet
a clear process is difficult to assess. They are correlated to
sea-level changes (Holland and Zimmermann 2000).
Hardie (1996) proposed that seawater chemical variations
in Phanerozoic were determined by the mixing ratio
between river water and its solution charge and mid-
ocean ridges solutions coupled with solid CaCO3 and
SiO2 phases. However, the model fails to explain the
near-constancy of the Kþ concentration that may be
related to the processes of potassium uptake by riverine
clays (Holland 2003). On the striking correlation between
the sea-level stand and seawater chemical variations,
Hardie (1996) suggested that the stand of the sea level
reflects the rate of oceanic crust formation (younger,
hotter, and rapid oceanic formation produces higher
sea levels). This, in turn, determines the rate of seawater
cycling through MOR and hence the mixing ratio of hydro-
thermally altered water versus river water. But one of the
most complete records of the seawater composition changes
is the curve of the Sr isotopic ratio (87Sr/86Sr) of Veizer et al.
(1989). This curve, refined through the years shows chem-
istry cycles that could be related to changes in the type of
rocks undergoing weathering during time (as also indi-
cated by the 187Os/186Os ratios) indicating the present
dominance of continental weathering as source of salinity
of the oceans.
Applications
A complete picture of the seawater geological history on
Earth could be extremely useful in determining the exis-
tence of past oceans on other telluric planets such as Mars,
through the study of “sedimentary” rocks and chemical
precipitates of “marine” origin.
Future Directions
The geological record of the seawater is still much incom-
plete and spotty. Bringing new evidences that pristine
seawater fluid inclusions could have been preserved
through eons and finding fluid inclusions containing
non-evaporated seawater are the most challenging
researches to carry out in the next decade to clearly asses
the chemical evolution of the oceans.
See also
▶ Archean Eon
▶ BIF
▶ Earth, Formation and Early Evolution
▶ Fluid Inclusions
▶ Fractionation, Mass Independent and Dependent
▶ Hadean
▶ Iron Oxyhydroxides
▶ Isua Supracrustal Belt
▶ Jack Hills (Yilgarn, Western Australia)
▶ Mid-Ocean Ridges
▶ Oceans, Origin of
▶ Oxygenation of the Earth’s Atmosphere
▶ Pillow Lava
▶ Siderite
▶ Snowball Earth
▶ Soda Lakes
▶ Sulfur Isotopes
▶ Zircon
References and Further Reading
Appel PWU, Rollinson HR, Touret JLR (2001) Remnants of an early
Archaean (> 3.75Ga) seafloor, hydrothermal system in the Isua
Greenstone Belt. Precambrian Res 112:27–49
Bickle MJ (1986) Implications of melting for stabilisation of the
litosphere and heat loss in the Archaean. Earth Planet Sci Lett
80:314–324
Cairns-Smith AG (1978) Precambrian solution photochemistry, inverse
segregation, and banded iron formations. Nature 276:807–808
Channer DMD, de Ronde CEJ, Spooner ETC (1997) The Cl-Br-I compo-
sition of 3.23 Ga modified seawater: implications for the geological
evolution of ocean halide chemistry. Earth Planet Sci Lett 150:
325–335
de Ronde CEJ, Channer DMD, Faure K, Bray CJ, Spooner TC (1997) Fluid
chemistry of Archean seafloor hydrothermal vents: implications for
the composition of circa 3.2 Ga seawater. Geochim Cosmochim Acta
61:4025–4042
DOE (1994) Handbook of methods for the analysis of the various
parameters of the carbon dioxide system in sea water. Version 2,
A. G. Dickson & C. Goyet, eds. ORNL/CDIAC-74
Farquhar J, Bao H, Thiemens M (2000) Atmospheric influence of Earth’s
earliest sulfur cycle. Science 289:756–758
Foriel J, Philippot P, Rey P, Somogyi A, Banks D, Menez B (2004) Biolog-
ical control of Cl/Br and low sulfate concentration in a 3.5-Gyr-old
seawater from North Pole, Western Australia. Earth Planet Sci Lett
228:451–463
Hardie LA (1996) Secular variation in seawater chemistry: an explanation
for the coupled variation in the mineralogies of marine limestones
and potash evaporates over the past 600 My. Geology 24:279–283
Holland HD (1972) The geologic history of seawater: an attempt to solve
the problem. Geochim Cosmochim Acta 36:637–651
Holland HD (1984) The chemical evolution of the atmosphere and
oceans. Princeton University Press, Princeton, 582 pp
Holland HD (2003) The geologic history of seawater. In: Elderfield H (ed)
Treatise on geochemistry, vol 6. Elsevier, Amsterdam, pp 583–625
Holland HD (2006) The oxygenation of the atmosphere and ocean. Trans
Roy Soc B 361:903–915

Holland HD, Zimmermann H (2000) The dolomite problem revisited.
Int Geol Rev 42:481–490
Holser WT (1963) Chemistry of brine inclusions in Permian salt from
Hutchinson, Kansas. In: Bersticker AC (ed) Symposium on salt
(first). Northern Ohio Geological Society, Cleveland, pp 86–95
Horita J, Zimmermann H, Holland HD (2002) The chemical evolution of
seawater during the Phanerozoic: implications from the record of
marine evaporites. Geochim Cosmochim Acta 66:3733–3756
Kempe S, Degens ET (1985) An early soda ocean? Chem Geol 53:95–108
Knauth LP (2005) Temperature and salinity history of the Precambrian
ocean: implications for the course of microbial evolution.
Palaeogeogr Palaeoclimatol Palaeoecol 219:53–69
Martin H, Claeys P, Gargaud M, Pinti D, Selsis F (2006) 6. Environmental
context. Earth Moon Planet 98:205–245
McCulloch MT, Bennett VC (1994) Progressive growth of the Earth’s
continental crust and depleted mantle: geochemical constraints.
Geochim Cosmochim Acta 58:4717–4738
Millero FJ (2003) Physicochemical controls on seawater. In: Elderfield H
(ed) Treatise on geochemistry, vol 6. Elsevier, Amsterdam, pp 1–21
Pinti DL (2005) The formation and evolution of the oceans. In: Gargaud M,
Barbier B, Martin H, Reisse J (eds) Lectures in astrobiology. Springer,
Berlin, pp 83–107
Sleep NH, Zahnle K, Neuhoff PS (2001) Initiation of clement surface
conditions on the earliest Earth. Proc Nat Acad Sci USA 98:3666–3672
Spencer RJ, Hardie LA (1990) Control of seawater composition by mixing of
river waters and mid-ocean ridge hydrothermal brines. In: Spencer RJ,
Chou I-M (eds) Fluid–mineral interactions: a tribute to H.P. Eugster.
Special Publication 2. Geochemical Society, San Antonio, pp 409–419
Veizer J, Hoefs J, Ridler RH, Jensen LS, Lowe DR (1989) Geochemistry of
Precambrian carbonates: I. Archean hydrothermal systems. Geochim
Cosmochim Acta 53:845–857
Veizer J, Ala D, Azmy K, Bruckschen P, Buhl D, Bruhn F, Carden GAF,
Diener A, Ebneth S, Goddéris Y, Jasper T, Korte C, Pawellek F,
Podlaha OG, Strauss H (1999) 87Sr/86Sr, d13C and d18O evolution
of Phanerozoic seawater. Chem Geol 161:59–88
Weiershauser L, Spooner E (2005) Seafloor hydrothermal fluids, Ben
Nevis area, Abitibi greenstone belt: implications for Archean
(2.7 Ga) seawater properties. Precambrian Res 138:89–123
Wilde SA, Valley JW, Peck WH, Graham CM (2001) Evidence from
detrital zircons for the existence of continental crust and oceans on
the Earth 4.4 Gyr ago. Nature 409:175–178
Ocean, Temperature of
▶ Precambrian Oceans, Temperature of
Oceanic Crust
Definition
Oceanic crust is the outer layer of the solid Earth beneath
the oceans. This ▶ crust is 6–9 km thick and comprises
3 main layers (from top to bottom): Layer 1, a thin layer
(typically less than 500 m) of deepwater unconsolidated
Oceans, Origin of O 1167
carbonate or siliceous sediment that disappear close to
the mid-ocean ridges; Layer 2A consisting of 0.5 km thick
glassy-to-finely crystalline basalt usually in the form of
pillow basalt and 1.5 km thick sheeted vertical diabase
dykes (Layer 2B); Layer 3 (5 km thick and of density
3.0 g cm3) made of gabbros (the intrusive form of basalt)
and ultramafic cumulates. The crust forms through partial
melting of upwelling mantle beneath mid-ocean spreading
centers; the layering results from differentiation of the
basaltic magma produced by this melting. On Earth,
the age of oceanic crust ranges from zero at spreading
centers to a maximum of about 180 Ma in the northwestern
Pacific.
See also
▶ Crust
▶ Obduction
▶ Oceans, Origin of
▶ Ophiolite
▶ Plate Tectonics
Oceans, Origin of
DANIELE L. PINTI1, NICHOLAS ARNDT2
1
GEOTOP & Département des Sciences de la Terre et de
l’Atmosphère, Université du Québec à Montréal, O
Montréal, QC, Canada
2
Maison des Géosciences LGCA, Université Joseph
Fourier, Grenoble, St-Martin d’Hères, France
Keywords
Comets, cool early Earth, D/H, Isua, Jack Hills zircons,
meteorites, volatiles, water
Definition
The term ocean applies to the entire body of saltwater
covering more than 70% of the Earth’s surface. From
a planetary point of view, the term can be extended to
the layer of liquid that partially or totally covers the surface
or pervades the subsurface of a planet or satellite. In
addition to water, the term includes other substances
such as ethane, methane, or ammonia that are in a liquid
state under the conditions of the planetary surface. An
analogue could be the hypothetical ocean of ▶ Titan. The
liquid body could be at the planetary surface in thermo-
dynamic equilibrium with an atmosphere or separated by
another layer. In the early history of the planets, it could be
1168 O Oceans, Origin of
an ocean of magma. Examples of subsurface oceans are
those predicted for Titan or beneath the smooth icy sur-
face of the satellites ▶ Europa and Enceladus. The origin of
the ocean entails the complex processes that led to the
formation of a stable body of liquid water in thermody-
namic equilibrium with an atmosphere on a planetary
body. The following overview focuses on the formation
of the terrestrial oceans.
Overview
There is good evidence that the oceans were present on the
▶ Earth in the early ▶ Archean (3.8 Ga ago) after the
▶ Late Heavy Bombardment and plausible evidence that
they were already present in the ▶ Hadean. The convinc-
ing evidence comes from the two oldest areas of volcanic
and sedimentary rocks – the ▶ Isua Supracrustal Belt,
West Greenland and ▶ Nuvvuagittuq, Quebec, Canada
(formally Porpoise Cove). The ages of the rocks in both
areas have been established at about 3.8 Ga (for the latter a
date of 4.3 Ga has been suggested; see ▶ Nuvvuagittuq). In
the Isua Supracrustal Belt, pillow basalts provide evidence
of underwater eruption, and in both belts, metasedimentary
rocks (▶ banded iron formations, metapelite, and ferru-
ginous quartzite) are the products of erosion, fluvial trans-
port, and subaqueous deposition. Although the setting of
some of these rocks may have been lakes or shallow seas, it
is far more probable that these rocks erupted or were
deposited in ocean basins.
The evidence for Hadean oceans is less direct, because
the geological record has been mostly wiped out by the
intense tectonic activity of the young Earth. Most evidence
comes from 4.4 to 4.2 Ga old detrital zircons (ZrSiO4)
from the Mt. Narryer quartzite and from ▶ Jack Hills
metaconglomerate, Western Australia (Mojzsis et al.
2001; Wilde et al. 2001). ▶ Zircon is a common U-rich
trace mineral in granitic magmas formed by melting of
a hydrous source. Parental rocks of the Jack Hills zircons
possibly formed from melting of subducted hydrated oce-
anic crust. The presence of such a crust requires interac-
tion between basaltic lava and seawater as is suggested by
the 18O/16O ratio (denoted as d18O) of the Jack Hills
zircons which extends to 7.4  0.7 ‰. This value can be
explained only by a mixing between the mantle source of
the zircons (d18O ca. 5.0 ‰) and a source enriched in 18O
compared to the mantle value and derived from
weathering and low-temperature alteration of oceanic
crust. The d18O values of Hadean, Archaean, and Prote-
rozoic zircons show a secular increase to values up to 10‰.
This can be explained by an increase of the amount of
supracrustal, high-d18O material available for melting and
assimilation, as the “wet” continental crust evolved and
matured during the Precambrian. The presence of d18O of
7.4  0.7‰ in a 4.4 Ga old detrital zircon of Jack Hills
could thus be the indirect and oldest record of interaction
of a magmatic source with liquid water at the surface of
the Earth (Peck et al. 2001). The combination of these
observations lead to the “▶ cool early Earth” hypothesis
according to which temperatures were clement, and
oceans existed, on the surface of the Earth as early as
4.4 Ga (Valley et al. 2002).
The source of the oceanic water is debated. It is com-
monly accepted that any primitive solar-type atmosphere
surrounding the Earth was rapidly blown off by UV and
X-rays produced by the sun during its proto-(T-Tauri)-
star stage. This primitive atmosphere probably did not
contain water because Earth accreted too close to the
Sun. Earth was composed of “dry” raw material related
to enstatite chondrite ▶ meteorites (Javoy 1997). The
present hydrosphere probably was derived from meteor-
itic and cometary material that was added after the moon-
forming impact (Morbidelli et al. 2000). Similarities
between the hydrogen isotopic ratio D/H of the whole
Earth (149 – 153  106), ocean water (155.7  106),
and the average D/H ratios of hydrated carbonaceous
chondrites (149  6  106) and Antarctic micrometeor-
ites (154  16  106) indicates that asteroids or asteroi-
dal dust are the most likely candidates as water carrier
(Maurette et al. 2000; Marty and Yokochi 2006).
▶ Comets apparently exhibit heavier D/H ratios of 290–
320  106 that are incompatible with oceanic water,
although only a few comets have measured D/H ratios.
Mass and isotopic balance calculations suggest that
a maximum of 10–20% of cometary water could have
contributed to the terrestrial water budget.
The total amount of water delivered to Earth was
certainly higher than the mass of the present-day oceans
(1.4  1021 kg), because the losses of volatiles to space
were important at the beginning. Pepin (1991) suggested
that at least 50% of the water delivered to Earth could have
dissociated to hydrogen and oxygen by the strong early UV
radiation. Further losses were due to impact erosion of the
atmosphere by arriving planetary bodies. Geophysical and
high-pressure mineralogy experiments suggest that the
modern mantle contains two to five times the ocean vol-
ume of water (Abe et al. 2000) and thus an amount of
water much higher than that preserved in the oceans was
added to the newly born Earth.
Theories of how the ocean formed and stabilized on
the surface of the Earth are based on geophysical modeling
and speculation. Water, together with CO2 (now trapped
at the Earth’s surface as carbonates), were vapor phases in
the secondary (outgassed) atmosphere. They did not

condense until the surface of the planet cooled down to
less than 647 K, the critical point of water. Assuming that
all oceanic water was present as a gas, the atmospheric
pressure was 270 bars of H2O plus 40 bars of CO2 (Sleep
et al. 2001; Pinti 2005). This runaway greenhouse atmo-
sphere, refurbished by a high heat flow from the still
molten interior of the Earth, kept the planet’s surface
sufficiently hot for several millions years to prevent water
from condensing on its surface. When a primordial crust
formed, it reduced the heat flux from the interior and the
surface of the Earth cooled down, allowing the condensa-
tion of water. It is difficult to assess when this occurred.
Geophysical models developed by Sleep et al. (2001) sug-
gest a few million years or less after the Moon-forming
impact, depending on the parameters chosen. If the geo-
chemistry of the Jack Hills zircons does indeed reveal the
presence of liquid water at the surface, this means that
between 4.4 and 4.3 Ga, oceans were already present, or, at
least, that liquid water was stable at the surface of the
Earth. The oceans became cold enough to assure the
survival of the first living communities sometime between
4.3 Ga and the end of the meteoritic bombardment at
3.85 Ga.
The chemistry of seawater in the early Hadean
was likely controlled by high-temperature fluid-rock reac-
tions between the hot CO2–H2O-rich atmosphere
and oceans, and primitive basaltic crust. Weathering of
Na-rich basalt reacting with chlorine saturated the seawa-
ter with NaCl. The acidity of water was possibly as high
as pH = 5 because the ocean was in thermodynamic
equilibrium with a CO2-rich atmosphere (Pinti 2005),
though alternative models of alkaline oceans saturated
with ▶ thermonatrite formed by weathering of the
basaltic protocrust have been proposed (Kempe and
Degens 1985).
Key Research Findings
To help understand the origin of the oceans requires the
integration of information from the Earth, where the
ocean still exists and where a fragmentary record of its
early state is preserved in the geological record, with infor-
mation from other planetary bodies. Studies of the Mar-
tian surface have already provided evidence of the small
shallow oceans that existed there early in the planet’s
history, and the cold icy satellites provide a record of
oceans of very different types. This information can
potentially be exported to models for the formation of
oceans on exoplanets. Léger et al. (2004), for example, has
proposed the existence of a hypothetical family of “Ocean-
Planets” that are entirely covered by an ocean hundreds of
kilometers deep.
Oceans, Origin of O 1169
Future Directions
Two main questions need to be addressed: (1) When did
oceans form on Earth? (2) What is the origin of the water?
The first question is difficult to answer because the
answers must be sought from a period when the geological
record is almost totally absent. Two decades ago the task
seemed hopeless, but the remarkable results emerging
from the Jack Hills zircons and other relicts of the oldest
crust have radically changed our knowledge of this period.
We need to search for and precisely date and analyze
other minerals or rocks that could provide complemen-
tary evidence of interaction with liquid water. For an
answer to the second question, we need to wait for future
space missions to study and collect samples from the
surface of comets (e.g., the Rosetta mission) to establish
to what extent this material could have contributed to the
terrestrial water budget.
See also
▶ Archea
▶ Banded Iron Formation
▶ Comet
▶ Cool Early Earth
▶ Degassing
▶ Earth, Formation and Early Evolution
▶ Europa
▶ Hadean
▶ Isotopic Fractionation (Interstellar Medium) O
▶ Isua Supracrustal Belt
▶ Jack Hills (Yilgarn, Western Australia)
▶ Late Heavy Bombardment
▶ Meteorites
▶ Nuvvuagittuq (Porpoise Cove) Greenstone Belt
▶ Rosetta (spacecraft)
▶ Soda Lakes
▶ Thermonatrite
▶ Titan
▶ Water, Delivery to Earth
▶ Zircon
References and Further Reading
Abe Y, Ohtani E, Okuchi T, Righter K, Drake MJ (2000) Water in the early
Earth. In: Righter K, Canup RM (eds) Origin of the earth and moon.
University of Arizona Press, Tucson, pp 413–433
Javoy M (1997) The major volatile elements of the Earth: their origin,
behavior, and fate. Geophys Res Lett 24:177–180
Kempe S, Degens ET (1985) An early soda ocean? Chem Geol 53:95–108
Léger A et al (2004) A new family of planets? “Ocean-Planets”. Icarus 169
(2):499–504
Marty B, Yokochi R (2006) Water in the early Earth. Rev Mineral
Geochem 62:421–450
1170 O Oceanus, Oceani

Maurette M, Duprat J, Engrand C, Gounelle M, Kurat G, Matrajt G,

Toppani A (2000) Accretion of neon, organics, CO2, nitrogen and
water from large interplanetary dust particles on the early Earth.
Planet Space Sci 48:1117–1137
Mojzsis SJ, Harrison MT, Pidgeon RT (2001) Oxygen-isotope evidence
from ancient zircons for liquid water at the Earth’s surface 4, 300 Myr
ago. Nature 409:178–181
Morbidelli A, Chambers J, Lunine JI, Petit JM, Robert F (2000) Source
regions and timescales for the delivery of water to the Earth. Meteorit
Planet Sci 35:1309–1320
Peck WH, Valley JW, Wilde SA, Graham CM (2001) Oxygen isotope ratios
and rare earth elements in 3.3–4.4 Ga zircons: ion microprobe
evidence for high d18O continental crust and oceans in the Early
Archean. Geochim Cosmochim Acta 65:4215–4229
Pepin RO (1991) On the origin and ealy evolution of terrestrial planetray
atmospheres and meteoritic volatiles. Icarus 92:2–79
Pinti DL (2005) The formation and evolution of the oceans. In: Gargaud
M, Barbier B, Martin H, Reisse J (eds) Lectures in astrobiology.
Springer, Berlin
Sleep NH, Zahnle K, Neuhoff PS (2001) Initiation of clement surface
conditions on the earliest Earth. Proc Natl Acad Sci USA
98:3666–3672
Valley JW, Peck WH, King EM, Wilde SA (2002) A cool early Earth.
Geology 30:351–354
Wilde SA, Valley JW, Peck WH, Graham CM (2001) Evidence from
detrital zircons for the existence of continental crust and oceans on
the Earth 4.4 Gyr ago. Nature 409:175–178
Oceanus, Oceani
Definition
The term Oceanus refers to a large low-albedo feature –
Oceanus Procellarum – on the ▶ Moon. Oceanus
Procellarum (ocean of storms) is the only surface feature
to which the term Oceanus is applied. It denotes the
largest ▶ Mare area on the Moon. Oceanus Procellarum
OGLE
▶ Optical Gravitational Lensing Experiment
OGLE-2005-BLG-390Lb
Definition
OGLE-2005-BLG-390Lb is a cool, ▶ super-Earth that was
discovered with the gravitational microlensing technique by
the Probing Lensing Anomalies NETwork/Robotic Telescope
Network (PLANET/Robonet), ▶ Optical Gravitational
Lensing Experiment (OGLE), and ▶ Microlensing Obser-
vations in Astrophysics (MOA) collaborations. The mass
and orbital separation of the planet are uncertain, as is the
mass and distance of the host star from the Sun, but the most
likely values are a mass of 5.5 Earth masses and projected
separation of 2.6 AU for the planet, and a mass of 0.22 solar
masses and a distance of 6.6 kpc for the host star. With an
equilibrium temperature of 50 K, this low-mass planet is
located in the cool, outer reaches of its planetary system; it is
the first such planet discovered by any technique.
The figure shows the observed light curve for the
OGLE-2005-BLG-390 microlensing event, including data
from the PLANET Danish, Perth, and Canopus telescopes,
RoboNet Faulkes North telescope, OGLE telescope, and
MOA telescope. The OGLE data over a wider span of time
is shown in the top left inset, whereas the planetary
3 3 1.6
OGLE
1.5
2
2.5 1.4
Magnification

Planetary
extends approximately 2,500 km across and covers 1 1.3 deviation
a surface area of 4 Mio. km2. It is composed of extensive 2 –1000 0 9.5 10 10.5 11
flood ▶ basalts but is not confined to an isolated impact
OGLE Danish
▶ crater or basin as it connects several ▶ Mare-type fea- 1.5 Robonet Perth
tures. Oceanus Procellarum was sampled during the Luna Canopus MOA

and Surveyor missions and was visited by the Apollo 12 1

astronauts in late 1969. –20 0 20
Days since 31.0 July 2005 UT
See also
▶ Albedo Feature OGLE-2005-BLG-390Lb. Figure 1 The figure shows the
▶ Apollo Mission observed light curve for the OGLE-2005-BLG-390 microlensing
▶ Basalt event, including data from the PLANET Danish, Perth, and
▶ Crater, Impact Canopus telescopes, RoboNet Faulkes North telescope, OGLE
▶ Impact Basin telescope, and MOA telescope. The OGLE data over a wider
▶ Mare, Maria span of time is shown in the top left inset, whereas the
▶ Moon, The planetary deviation is highlighted in the top right inset
▶ Palus, Paludes (Beaulieu et al. 2006).
 OGLE-2006-BLG-109Lb,c O 1171

deviation is highlighted in the top right inset (Beaulieu Is was discovered by the ▶ Microlensing Follow-Up NET-
et al. 2006). work (MicroFUN) collaboration, in conjunction with the
▶ Optical Gravitational Lensing Experiment (OGLE),
See also ▶ Microlensing Observations in Astrophysics (MOA),
▶ Exoplanets Discovery
and Probing Lensing Anomalies NETwork/Robotic Tele-
▶ Microlensing Observations in Astrophysics (MOA)
scope Network (PLANET/Robonet) collaborations. This
▶ Microlensing Planets
system consists of two planets with masses similar to
▶ Optical Gravitational Lensing Experiment
Jupiter and Saturn orbiting a star with roughly half the
▶ Probing Lensing Anomalies Network
mass of the sun located roughly 1.5 kPc from the Earth.
This system is analogous to our Jupiter/Saturn system, in
References and Further Reading that the two planets have the same mass ratios as Jupiter
Beaulieu JP et al (2006) Discovery of a cool planet of 5.5 Earth masses
and Saturn as well as the same ratio of orbital radii, and
through gravitational microlensing. Nature 439:437–440
the same (inferred) equilibrium temperatures (Gaudi et al.
2008; Bennett et al. 2010).
The Fig. 1 shows the observed light curve for the
OGLE-2006-BLG-109Lb,c OGLE-2006-BLG-109 microlensing event, including data
from OGLE, various MicroFUN telescopes, MOA,
Definition PLANET, and RoboNet. Five features are labeled. Four of
OGLE-2006-BLG-109Lb,c is the first multi-planet system these are due to the Saturn-mass planet, whereas the fifth
discovered with the gravitational microlensing technique. can only be explained by a second, Jupiter-mass planet.

A 3 OGLE
mFUN Auckland
4 mFUN Wise
14
mFUN MDM
mFUN CTIO I
mFUN CTIO H O
1 mFUN Mt. Lemmon
2 mFUN New Mexico
0.01q E mFUN Farm Cove
MOA
I magnitude

PLANET Canopus
15 2 RoboNet La Palma

5
3 5
B 4

16 1

3820 3825 3830 3835

HJD–2450000.

OGLE-2006-BLG-109Lb,c. Figure 1 The figure shows the observed light curve for the OGLE-2006-BLG-109 microlensing
event, including data from OGLE, various MicroFUN telescopes, MOA, PLANET, and RoboNet. Five features are labelled. Four of
these are due to the Saturn-mass planet, whereas the fifth can only be explained by a second, Jupiter-mass planet. The inset
shows the path of the source through the caustic, with the locations of the source which give rise to the five features labelled.
1172 O OH

The inset shows the path of the source through the caustic,
with the locations of the source which give rise to the five
features labeled.
See also
▶ Exoplanets Discovery
▶ Microlensing Follow-Up Network
▶ Microlensing Observations in Astrophysics
▶ Microlensing Planets
▶ Optical Gravitational Lensing Experiment
▶ Probing Lensing Anomalies Network
References and Further Reading
Bennett DP et al (2010) Astrophys J 713:837
Gaudi BS et al (2008) Discovery of a Jupiter/Saturn Analog with gravita-
tional microlensing. Science 319(5865):927
Oligomer
Definition
The term oligomer is derived from the Greek “oligos,”
meaning “a few.” In chemistry, an oligomer is a short
polymer consisting of approximately five monomer
units, although agreement as to the strict length cutoff
is debated and varies between four and one hundred.
Oligomers are formed by oligomerization, which involves
linking the monomer units together in a chemical reac-
tion. Unlike a ▶ polymer, if one of the repeating units of
oligomer is removed, its chemical properties may be sig-
nificantly altered.
In biochemistry, the term oligomer is commonly used
for short, single-stranded DNA fragments, used in
▶ hybridization experiments as ▶ primers. It can also
indicate a protein made of two or more subunits.

See also
▶ Hybridization
▶ Nucleic Acids
OH ▶ Polymer
▶ Primer
▶ Hydroxyl Radical ▶ Protein

Oligomerization
Oligarchic Growth
Definition
Definition Oligomerization is a chemical process that links mono-
Oligarchic growth is the second-to-last stage of the for- meric compounds (e.g., amino acids, nucleotides, or
mation of terrestrial planets and giant planet cores. During monosaccharides) to form dimers, trimers, tetramers, or
this stage, the largest planetary embryos grow quickly longer chain molecules (oligomers). Examples are the
while the smallest grow slowly, leading to a bifurcated conversion of nucleotides to oligonucleotides and amino
mass distribution with a number of lunar- to Mars-mass acids to peptides. The boundary between what is consid-
embryos embedded in a swarm of smaller planetesimals. ered an oligomer and a ▶ polymer is unclear, but it is
Oligarchic growth transitions to chaotic growth when the usually accepted to be in the range of 10–100 monomer
planetary embryos become large enough to overcome units. Prebiotic experiments have shown that ▶ activated
eccentricity damping due to dynamical friction from nucleotides can be oligomerized using ▶ clay minerals
smaller planetesimals. This leads to embryo–embryo col- as catalyst.
lisions and signals the last stage of accretion of terrestrial
planets and giant planet cores. See also
▶ Activated Nucleotide
▶ Amino Acid
See also ▶ Clay
▶ Late-Stage Accretion ▶ Mineral
▶ Planetary Formation ▶ Monosaccharide
▶ Planetesimals ▶ Nucleotide
▶ Runaway Growth ▶ Oligonucleotide

▶ Oligopeptide
▶ Polymer
▶ Prebiotic Chemistry
Oligonucleotide
KUNIO KAWAMURA
Department of Applied Chemistry, Osaka Prefecture
University, Naka-ku, Sakai, Osaka, Japan
Synonyms
DNA; Nucleotide oligomer; RNA
Keywords
Chemical evolution, nucleotide, oligonucleotide, RNA
Definition
An oligonucleotide is a relatively short RNA or DNA
polymer linked via phosphodiester bonds. Under prebi-
otic conditions, oligonucleotides of RNA can be formed
from ▶ activated nucleotide monomers including different
kinds of nucleotide bases: ▶ guanine, ▶ adenine, ▶ uracil,
▶ cytosine, and ▶ hypoxanthine. The oligonucleotides
of RNA formed under the primitive earth conditions
normally contain both 20 ,50 - or 30 ,50 - linkages and often
contain a 50 -residue linked by a diphosphate linkage.
Overview
The formation of oligonucleotides would have been an
essential step for the emergence of life on the primitive
earth. In modern organisms, polymerization of RNA or
DNA monomers merely proceeds from nucleoside
50 -triphosphate monomers in the presence of DNA tem-
plate and specific enzymes. On the contrary, the prebiotic
formation of RNA with 50 nucleotide units in length
proceeds from nucleoside 50 -phosphorimidazolides
under simulated primitive earth conditions. However,
the prebiotic formation of DNA does not proceed from
such activated nucleotide monomers.
Although the usage of condensation reagents is possi-
ble to form oligonucleotides from nucleotide monomers,
the activated nucleotides are more effectively used to form
long oligonucleotides with high yield. Continuous inves-
tigations have been carried out to identify the pathways for
prebiotic formation of RNA oligonucleotide. Successful
examples include the oligonucleotide formation from the
activated nucleotide monomers in the presence of metal
Oligonucleotide O 1173
ions (Sawai 1976), that in the presence of montmorillonite
clay catalyst (Ferris and Ertem 1992; Huang and Ferris
2006), that in the presence of a polynucleotide template
(Inoue and Orgel 1983). Normally, the condensation of the
activated nucleotide monomers provides oligonucleotides
with an average length of 10 units in the presence of metal
ion catalyst or clay mineral catalyst. Besides, spontaneous
condensation of the activated nucleotide by continuous
addition of the activated nucleotide in the presence of
▶ montmorillonite clay results in the formation of oligo-
nucleotides 50-mer (Ferris et al. 1996). On the other
hand, oligoguanylates up to 40-mer in length form by
the condensation of guanosine 50 -phosphorimidazolide
on a polycytidylate template with Zn2+ ion, where the
Watson–Crick hydrogen bonding directs the activated
monomers of guanosine nucleotide on the template poly-
nucleotide. However, this template-directed reaction is
not efficient for the condensation of adenosine, uridine,
or cytidine 50 -phosphorimidazolide on the complemen-
tary polynucleotide template. This is probably due to the
fact that the template-directed formation of oligonucleo-
tides is mainly supported by the stacking interactions
between the activated nucleotide monomers rather than
the Watson–Crick hydrogen bonding between the acti-
vated nucleotide and the complementary template. At the
same time, it has been confirmed that the replication of
oligonucleotides partially proceeds from the mixture of
four types of activated nucleotide monomers in the
presence of mixed base template polynucleotides. O
See also
▶ Activated Nucleotide
▶ Montmorilllonite
▶ Nucleic Acids
▶ Nucleoside Phosphoimidazolide
▶ RNA World
▶ Self Replication
▶ Template-Directed Polymerization
References and Further Reading
Ferris JP, Ertem G (1992) Oligomerization of ribonucleotides on mont-
morillonite: reaction of the 50 -phosphorimidazolide of adenosine.
Science 257:1387–1389
Ferris JP, Hill Jr AR, Liu R, Orgel LE (1996) Synthesis of long prebiotic
oligomers on mineral surfaces. Nature 381:59–61
Huang W, Ferris JP (2006) One-step, regioselective synthesis of up to
50-mers of RNA oligomers by montmorillonite catalysis. J Am Chem
Soc 128:8914–8919
Inoue T, Orgel LE (1983) A nonenzymatic RNA polymerase model.
Science 219:859–862
Sawai H (1976) Catalysis of internucleotide bond formation by divalent
metal ions. J Am Chem Soc 98:7037–7039
1174 O Oligopeptide
Oligopeptide
JEAN-FRANÇOIS LAMBERT
Laboratoire de Réactivité de Surface, Université Pierre et
Marie Curie, Paris, France
Synonyms
Peptide
Keywords
Activation, amino acids, peptides, proteins
Definition
An oligopeptide is a short-chain ▶ peptide, i.e., a polymer
of ▶ amino acids (AAs) connected by amide, or more
precisely peptide, linkages. The term is usually limited to
peptides with less than 20–25 amino acid residues.
Overview
In “peptides-first” scenarios for the emergence of biomacro-
molecules, oligopeptides capable of specific catalysis of some
biological reactions (primitive metabolism) are suggested to
have appeared without the previous existence of genetic
material. For instance, in his protometabolic model, de
Duve (1991) calls these short peptides, “multimers.”
Chemically, the emergence of oligopeptides from
preexisting amino acids poses several problems. The first
two are consequences of chemical thermodynamics. The
condensation of an amide bond between two AAs is ther-
modynamically unfavorable in solution, thus the amount
of oligopeptide in an amino acid solution at equilibrium
falls quickly with the peptide chain length: for instance,
a glycine solution with a total AA concentration of 0.5 M,
at equilibrium, would contain only one molecule of
18-mer per hundred cubic kilometers!
In addition, the formation of oligomers from
a mixture of different amino acids would likely show little
selectivity, i.e., all possible sequences would be randomly
formed. Since the number of sequences increases
exponentially with the peptide length, the probability of
forming significant amounts of a specific oligomer is van-
ishingly low.
The last problem concerns the slow kinetics of oligo-
merization: at room temperature and moderate pH, com-
pletion of the reaction would take several centuries due to
high activation energy barriers.
Several solutions have been proposed to allow the
prebiotic emergence of oligopeptides with a reasonable
probability:
– They could have formed in submarine hydrothermal
vents: thermodynamic equilibria are displaced toward
AA oligomerization in hydrothermal conditions,
although probably not enough to give high peptide
yields.
– Oligomerization could have been coupled to
a thermodynamically downhill chemical process.
This would involve the input of “high-energy mole-
cules” (more properly, molecules with high free
enthalpies of formation) such as nitrogen oxides.
– Alternatively, it may have occurred in the “adsorbed
state,” i.e., after adsorption of the AAs on mineral
surfaces (“polymerisation on the rocks,” Orgel 1998).
This hypothesis connects well to “surface metabolism”
scenarios (e.g., those of Wächtershäuser) and may
solve the kinetic as well as the thermodynamic prob-
lem, since catalysis by surface groups is likely.
The last two scenarios have been shown empirically to
result in the formation of oligopeptides up to the
pentamers at least, even though molecular details remain
sketchy. It is noteworthy that they are only successful in
fluctuating environments, i.e., when coupled with varia-
tions of the macroscopic conditions such as wetting and
drying cycles.
See also
▶ Amino Acid
▶ Peptide
▶ Polypeptide
▶ Oligonucleotide
References and Further Reading
Brack A (2007) From interstellar amino acids to prebiotic catalytic pep-
tides: a review. Chem Biodivers 4:665–679
Cleaves HJ, Aubrey AD, Bada JL (2009) An evaluation of the critical
parameters for abiotic peptide synthesis in submarine hydrothermal
systems. Orig Life Evol Biosph 39:3109–3126
Commeyras A, Taillades J, Collet H, Boiteau L, Vandenabeele-Trambouze O,
Pascal R, Rousset A, Garrel L, Rossi J-C, Biron J-P, Lagrille O,
Plasson R, Souaid E, Danger G, Selsis F, Dobrijévic M, Martin H
(2004) Dynamic co-evolution of peptides and chemical energetics,
a gateway to the emergence of homochirality and the catalytic activity
of peptides. Orig Life Evol Biosph 34:35–55
de Duve C (1991) Blueprint for a cell: the nature and origin of life. Neil
Patterson, Burlington, NC
Lambert J-F (2008) Adsorption and polymerization of amino
acids on mineral surfaces: a review. Orig Life Evol Biosph 38:211–242
Leslie E (1998) Orgel polymerization on the rocks: theoretical
introduction. Ori Life Evol Biosph 28(3):227–234
Rode BM (1999) Peptides and the origin of life. Peptides 20:773–786
Zaia DAM (2004) A review of adsorption of amino acids
on minerals: was it important for origin of life? Amino Acids
27:113–118

Oligosaccharide
▶ Carbohydrate
Olympus Mons
Definition
Olympus Mons is the largest known ▶ volcano on ▶ Mars
and in the entire ▶ Solar System (height above the Martian
topographic datum: 22 km; height above surrounding
plains: 20 km; basal diameter: 640 km). The flanks
are covered by tube-fed and channel-fed lava flows and
have slopes of typically 3–5
, except for a steeper, up to
6 km-high basal scarp. The morphological analogy to
terrestrial volcanoes (e.g., Hawaii, Galapagos) suggests
that Olympus Mons is a basaltic shield volcano. A summit
caldera has 80 km diameter and developed in multiple
stages over the last several hundred million years (Ma).
Some lava flows are probably even younger (10 Ma),
indicating possible ongoing volcanism on Mars.
See also
▶ Basalt
▶ Mars
▶ Solar System, Inner
▶ Tharsis
▶ Volcano
Oort Cloud
JACQUES CROVISIER
LESIA - Bâtiment ISO (n
17), Observatoire de Paris,
Meudon, France
Keywords
Comets, small bodies
Definition
The Oort cloud is a spherical cloud of ▶ comets surround-
ing the Solar System and extending out to heliocentric
distances greater than 100,000 AU.
Overview
The idea of a distant reservoir of ▶ comets, extending to
the verge of the ▶ Solar System, was already discussed by
Oort Cloud O 1175
Giovanni Schiaparelli (1910) and Ernst Öpik (1932). The
Dutch astronomer Jan Oort (1900–1992) pointed out in
1950 the need of such a reservoir to explain the continuous
input of new long-period comets. He showed, from the
data gathered by Van Woerkom (1948) that many long-
period comets have their aphelia at more than 20,000 AU.
This reservoir is now known as the “Oort cloud.”
The Oort cloud is supposed to be spherical, extending
from 20,000 ▶ AU from the Sun to 200,000 AU (the limit
of the gravitational influence of the Sun). It could com-
prise as many as 1012 comets.
These comets cannot be directly observed in the Oort
cloud, being too far from the Sun to manifest any activity.
Following perturbations by a nearby star (or by a hypo-
thetical distant planet), they may be transferred to orbits
bringing them close to the Sun, where they become active.
Such “Oort-cloud comets” have random inclination
over the ecliptic and belong to the family of “nearly-
ISOTROPIC comets.” The most famous of them is 1P/
▶ Halley.
Comets stored in the Oort cloud were not formed
there. They were formed in the inner Solar System (pre-
sumably in the Jupiter – Uranus region) and subsequently
ejected to the outer Solar System following gravitational
perturbations by the giant planets.
The Oort cloud contrasts with the alternate reservoir
of comets, the ▶ Kuiper belt, which is at the origin of
Jupiter-family comets.
O
See also
▶ Comet
▶ Kuiper Belt
▶ Trans-Neptunian Object
References and Further Reading
Barucci MA, Boehnhardt H, Cruikshank DP, Morbidelli A (2008)
The solar system beyond Neptune. University of Arizona Press,
Tucson
Dones L, Weissman PR, Levison HF, Duncan MJ (2005) Oort Cloud
formation and dynamics. In: Festou MC, Keller HU, Weaver H
(eds) Comets II. University of Arizona Press, Tucson, pp 153–174
Festou MC, Keller HU, Weaver H (2005) Comets II. University of Arizona
Press, Tucson
Oort JH (1950) The structure of the cloud of comets surrounding the
Solar System, and a hypothesis concerning its origin. Bull Astron Inst
Neth 11:91–110
Öpik EJ (1932) Note on stellar perturbations of nearly parabolic orbits.
Proc Am Acad Arts Sci 67:169–183
Schiaparelli G (1910) Orbites cométaires, courants cosmiques,
mŽtŽorites. Bull Astron Ser I 27:194–205
Van Woerkom AJJ (1948) On the origin of comets. Bull Astron Inst Neth
10:445–472
1176 O Opacity
Opacity
Synonyms
Attenuation
Definition
Opacity measures the property of a medium to attenuate
light. The opacity depends on the composition of the
medium, its density, and temperature, but also on the
wavelength considered. The absorption coefficient an
(with units [cm1]) enters into the definition of the
▶ optical depth. More often in astronomy, it is the mass
absorption coefficient kn ¼ an =r (with units [cm2 g1])
which is used.
The main sources of opacity in an astronomical
context are:
● Electron scattering (at temperatures of the order
1 billion K)
● Electronic transitions (free-free, bound-free, or
bound-bound at temperature of the order 10,000 K)
● Molecular or dust absorption (around or below
3,000 K)
The opacity plays an essential role in radiative transfer,
and thus in the energetic equilibrium of stars.
See also
▶ Mean Free Path
▶ Optical Depth
Opaline Silica
Definition
Opaline silica (SiO2 · nH2O) occurs at several locations on
▶ Mars, although the definitive identification is conten-
tious. The confirmed finding would indicate past aqueous
activity. Aqueous free silica is a product of ▶ basalt
weathering, when the interaction of water with mafic
(i.e., Mg- and Fe-rich, silica-poor) ▶ rock rapidly dissolves
olivine, pyroxene, and glass. Opaline silica could have been
precipitated in hydrothermal or lacustrine (related to
a lake) evaporitic environments on Mars. On Earth, it rarely
persists more than a few million years, because it rapidly
transforms into microcrystalline quartz during ▶ diagene-
sis. The persistence of opaline silica on Mars over much
longer geologic timescales (billions of years) indicates that
water was not present for extended periods of time.
See also
▶ Basalt
▶ Chert
▶ Diagenesis
▶ Hydrothermal Environments
▶ Mars
▶ Rock
▶ Weathering
Oparin’s Conception of
Origins of Life
STÉPHANE TIRARD
Faculté des Sciences et des Techniques de Nantes, Centre
François Viète d’Histoire des Sciences et des Techniques,
EA 1161, Nantes, France
History
In 1924, Alexander Ivanovitch Oparin (1894–1980),
a young scientist in vegetal physiology, published in Mos-
cow a very important text about the origin of life. With
this text, he began a very long career devoted to this
topic. He wrote important books and developed very
active experimental researches.
He published this first text, entitled Origins of Life, in
Russian. He described the evolution of earth and the
transformation of matter from mineral molecules to
organic molecules. Finally, according to him, little drops
of organic matter could appear, which constituted the last
step before becoming cells.
In 1936, Oparin published an important book entitled
Origins of Life. In an interdisciplinary approach, he
suggested a broad scenario of the origins of life on Earth,
describing all the steps of the evolution of the Earth, of
matter, and of primordial life. This book knew a great
success and was translated into several languages, and
especially into English in 1938.
This book is an exhaustive presentation of Oparin’s
ideas, very close to his 1924 conception. However, he
introduced a lot of current scientific data and two specific
points have to be underlined. Firstly, Oparin introduced
the notion of coacervate coming from Bungenberg de
Jong’s work. Coacervates were microscopical vesicles
formed in colloidal solutions and Oparin used them as
a model of a primitive cell in his theory. Secondly, Oparin
claimed that the primitive atmosphere did not contain
CO2, and that it was reductive.

After the Second War, this book still was a point of
reference. For example, John Desmond Bernal (1951)
quoted it. In 1952, ▶ Harold Urey, the American physicist,
wrote a synthesis on the primitive conditions and referred
to Oparin (1938) about the lack of CO2 in the primitive
atmosphere and agreed with the Soviet scientist on this
point.
During the following three decades, Oparin, who had
important academic positions in the USSR, continued to
be the leader of this topic in his country. He particularly
worked on primitive metabolism and developed a hetero-
trophic hypothesis on the origin of life on Earth. However,
in the lyssenkoist context, he neglected to work on hered-
ity molecules.
See also
▶ Bernal’s Conception of Origins of Life
▶ Calvin’s Conception of Origins of Life
▶ Haldane’s Conception of Origins of Life
▶ Miller, Stanley
▶ Monod’s Conception of Origins of Life
▶ Urey’s Conception of Origins of Life
References and Further Reading
Deamer DW, Fleischaker GR (1994) Origins of life, the central concept.
Jones and Bartlett, Boston
Oparin AI (1938) The origin of life, Sergius Morgulis trans. Macmillan,
New York
Open Cluster
Definition
An open star cluster is a loosely bound group of a few 103
stars formed from the same giant molecular cloud and
thus having the same age and ▶ metallicity (chemical
composition). This property makes them valuable tools
for the study of stellar evolution. Contrary to ▶ globular
clusters, open clusters are relatively young (less than a few
108 years old) and they are found in the plane of the Milky
Way. About a thousand open clusters are known in the
Galaxy, but their total number may be ten times higher.
The best known example is the Pleiades (108 year). Various
processes (tidal interactions with giant molecular clouds
and spiral arms) tend to disperse its members and dissolve
an open cluster.
See also
▶ Globular Cluster
▶ Metallicity
Ophiolite O 1177
Operational Taxonomic Unit
▶ Phylotype
Operon
Synonyms
Polycistronic transcript
Definition
An operon is a group of ▶ genes whose products have
related or complementary functions and which are tran-
scribed as a unit. Polycistronic or multi-gene are then
independently translated by ribosomes into the individ-
ual, functionally related proteins. Many bacterial and
archaeal genes are organized into operons, while they are
exceptional in eukaryotes. The operon model was pro-
posed by F. Jacob and J. Monod in 1961 to explain the
coordinated regulation of co-transcribed genes involved in
the metabolism of lactose in Escherichia coli. The so-called
Lac operon spans about 6,000 nts and includes a promoter,
an operator, three adjacent structural genes coding for
the enzymes required for lactose metabolism, and a termi-
nator. Regulation of the Lac operon was the first complex
genetic regulatory mechanism to be elucidated. O
See also
▶ Archea
▶ Bacteria
▶ Gene
▶ Genome
▶ Transcription
Ophiolite
Definition
An ophiolite is a slice of ▶ oceanic crust and underlying
mantle that is thrust onto a continent by ▶ obduction.
The stratigraphic sequence observed in an ophiolite cor-
responds to layered oceanic crust: from top to bottom, an
upper layer of oceanic sediment (siliceous or carbonate
ooze); a layer of pillow basalt; a layer of sheeted near-
vertical dykes; layers of ▶ gabbro and plagioclase and
mafic cumulates; and a basal layer of “tectonite,” the
1178 O Optical Depth

deformed harzburgite (a type of ▶ peridotite) of the

uppermost mantle. Many ophiolites contain only some
of these lithologies. Debate surrounds the question of
whether any ophiolite represents normal oceanic crust
formed at a mid-oceanic spreading centre. The geochem-
ical compositions of basalt rocks indicate that many
ophiolites are segments of crust that formed in back-arc
basins. Reactions between water and ophiolite at high
temperatures (300
C) can produce low-molecular hydro-
carbons by abiotic Fischer-Tropsch-Type mechanisms
during serpentinization.
See also
▶ Fischer-Tropsch-Type Reaction
▶ Gabbro
▶ Obduction
▶ Oceanic Crust
▶ Peridotite
▶ Pillow Lava
▶ Plate Tectonics
Optical Gravitational Lensing
Experiment
Synonyms
OGLE
Definition
OGLE is a collaboration of primarily Polish astronomers
that uses wide-field temporal monitoring to study a broad
range of astrophysical phenomena. The original goal of the
OGLE collaboration was to search for dark matter with the
gravitational microlensing technique. However, since its
inception, OGLE has also played a central role in the search
for ▶ exoplanets with microlensing. In addition, OGLE
identified the first transiting planets found by photometry
(as opposed to radial-velocity planets), and has contributed
to the study of variable stars, Galactic structure, strong
gravitational lenses, and Kuiper belt objects. The Principal
Investigator of the OGLE collaboration is Andrej Udalski
from Warsaw University. The OGLE 1.3-m telescope is
located at the Las Campanas Observatory in Chile.

See also
▶ Exoplanets Discovery
Optical Depth ▶ Microlensing Planets

Synonyms
Optical thickness
Optical Thickness
Definition
The optical depth is a dimensionless quantity, ▶ Optical Depth
generally noted t, that measures how absorbing – or ▶ scat-
tering – a slab in a medium is. Depending on whether
t < 1 or t > 1, a slab of medium is said to be optically thin
or optically thick. In a medium of constant density,
the optical depth is proportional to the thickness of Optical Rotatory Power
the slab.
In an absorbing medium, the intensity of inward radi- ▶ Rotatory Power
ation decreases as exp(t). In an emitting medium, such
as a stellar photosphere, or a planetary atmosphere in the
infrared, the outward radiation is well approximated by
assuming that it comes entirely from the layer of optical
depth t = 1.
Orbit
AVI M. MANDELL
See also NASA Goddard Space Flight Center, Greenbelt, MD, USA
▶ Extinction, Interstellar or Atmospheric
▶ Opacity
▶ Radiative Transfer Keywords
▶ Scattering Gravity, Kepler, Newton

Definition
The orbit of a celestial body is its path around a central
mass, caused by the acceleration due to the gravitational
force between them. The planets and minor bodies such as
asteroids and comets in the Solar System orbit the Sun,
while any celestial body in the Solar System that orbits
something other than the Sun is called a satellite (moon).
Long-term orbital motion is affected by the gravitational
forces due to all bodies in the vicinity, as well as to torques
due to forces such as gas drag, dynamical friction, and
radiation pressure.
History
The concept of a celestial orbit can be generalized to
include any motion of a celestial body around another
body, and dates back at least to the earliest written texts.
However, Johannes Kepler was the first to establish the
true qualitative characteristics of astronomical orbits, for-
mulating his three laws of planetary motion between 1609
and 1619. Analyzing the results from painstaking obser-
vations of the motions of astronomical bodies by Tycho
Brahe over many years, he determined that these bodies
travel on elliptical orbits (with the Sun at one focus),
speeding up as they approach the Sun and then slowing
down as they move farther away. As simple as they seem
today, these two conceptual breakthroughs revolutionized
the Western world’s view of the structure of the universe.
Until this time, the majority view was that the Sun,
planets, and star all revolved around the Earth; this theory
is known as geocentrism, with the most well-known pro-
ponent being the second-century Greco-Roman philoso-
pher Ptolemy. Heliocentrism, the idea that the orbits of
the planets are centered on the Sun, had first been pro-
posed by Aristarchos of Samos in the third century BCE,
but was largely ignored until it was explored by Coperni-
cus in 1543 and later supported by Galileo’s observations
of the motions of Jupiter’s moons. However, these early
models conceived the orbits as perfect circles, and there-
fore required many ad hoc additions to fit the observed
motions of planets in the sky. Kepler’s realization that the
orbits could be elliptical turned heliocentrism into a viable
predictive theory.
Kepler’s third law, that the square of the period of
a planet’s orbit is proportional to the cube of the distance
from the Sun (P2 / a3), provided the first mathematical
relation between two orbital quantities and formed one of
the primary drivers for Isaac Newton’s derivation of the
inverse square law of universal ▶ gravitation (Fgrav / 1/r2).
Newton was able to show that two gravitationally bound
bodies will orbit the center of mass of the two-body
system, and if the difference in mass is large enough the
Orbit O 1179
smaller body will essentially orbit the more massive body.
This simple mathematical formulation (with modifica-
tions included for general relatively as proposed by Albert
Einstein) forms the backbone of all of celestial mechanics
and dynamical astronomy.
Overview
As first envisioned by Newton, orbital shapes can be orga-
nized into three different categories based on how much
energy the orbiting body has: elliptical (including circu-
lar), parabolic, and hyperbolic. Parabolic and hyperbolic
orbits are unbound, in that they begin and end at an
infinite distance from the center of mass; the difference
between the two is that a body on a parabolic orbit has
zero net energy, while a hyperbolic orbit requires a positive
total energy and therefore has a nonzero beginning and
ending velocity. Newton realized that these shapes corre-
spond to sections of a cone (conic sections), with one
focus of the orbit lying on the rotational axis of the cone.
The conic section therefore confines the orbital motion to
a two-dimensional plane, known as the orbital plane.
Basic Methodology
It is important to start with a discussion of terminology,
since the language of celestial mechanics (the name for the
study of orbital dynamics) can become very confusing due
to the many specialized terms used to describe the prop-
erties of an orbit.
The general terms for the closest and farthest points of O
an orbit are the periapsis and apoaspsis; these terms can be
specialized for individual situations (i.e., perigee for orbits
around the Earth, perihelion for orbits around the Sun,
and periastron for orbits around any star). The line that
connects the periapsis and the apoapsis is called the line of
apsides; this line naturally passes through the central body,
and the direction of the periapsis can rotate to trace
a circle if the orbit is perturbed (known as apsidal
precession).
The motion of an orbiting body can be described by six
quantities known as orbital elements. These quantities can
be coordinates of position and velocity, such as the
Cartesian coordinates of a body in a specific reference
frame; but more commonly orbits are described using
the Keplerian orbital elements, since these are independent
of a specific reference direction. The six Keplerian elements
are the semimajor axis and eccentricity of the ellipse, and
the inclination, argument of periapsis, longitude of the
ascending node, and either the mean (temporal) or true
(angular) anomaly (see Fig. 1). The most useful and
general elements are semimajor axis, eccentricity, and
inclination, which describe the size, shape, and tilt of the
1180 O Orbit
Celestial body
True anomaly
ν
Argument of periapsis
ω
Ω Applications
γ
Longitude of ascending node Reference
direction
Plane of
referenc i deduce the orbital motions and positions of bodies in
e
Inclination
Ω our own system by studying the proper motion of objects
Ascending node
it
Orb
Orbit. Figure 1 A diagram of an orbit, with several orbital
elements labeled (inclination, argument of periapsis,
longitude of the ascending node, and true anomaly). The
additional orbital elements not labeled are the semimajor axis
(i.e., size of the orbit) and the eccentricity (i.e., ellipticity) of the
orbit (WikiCommons)
orbit with respect to the reference orbital plane (the plane
of reference in the Solar System is known as the ecliptic
plane). The semimajor axis is defined as one half of the
major axis of the orbital ellipse; note that this is different
than the distance to the central body when the orbit is not
a circle. The eccentricity describes how elliptical the orbit
is, ranging from 0 for a circular orbit to 1 for a parabolic
orbit. The inclination of the orbit is measured in degrees,
ranging from 0
to 180
(values larger than 90
indicate a
“retrograde” orbit that moves in the opposite direction
around the star). The other elements are less intuitive and
less frequently used, since they primarily define a single
orbital position and average out to zero for a population of
randomly oriented bodies (as opposed to eccentricity and
inclination that have a characteristic distribution for
a randomized population), and they will not be discussed
in detail here.
There are a number of processes that can lead to an
evolution of the orbital parameters of a single body or
a population of bodies; these effects are known as pertur-
bations. These perturbations generally fall into two cate-
gories: gravitational perturbations due to interactions with
other bodies in the system (such as secular perturbations
or dynamical friction), or relatively steady-state forces that
either add or remove energy from the orbit (such as gas
drag or radiation pressure). Gravitational forces between
two bodies will lead to an excitation of the orbit of the
smaller body (such as the scattering of comet-sized bodies
into the Oort cloud by Jupiter) and a damping of the
orbital excitation of the larger body (such as the damping
of planetary eccentricities during ▶ planet formation due
to interactions with small planetesimals).
The study of orbital dynamics is fundamental to much of
planetary astronomy in our own Solar System, as well as
the study of other stellar and planetary systems. We
as they move across the sky over time – the same method
has been used since classical times, and we are still using it
to discover ▶ Kuiper Belt objects at the far reaches of the
Solar System today. In other planetary systems, we can
measure the radial velocity of the central star and use
that motion to calculate the orbital motion of surround-
ing planets even when they are impossible to detect
directly.
We can also explore the evolution of the orbits of
bodies in planetary systems over time using computer
simulations. Computational numerical simulations of
planetary dynamics all aim to solve the same general
problem: how to calculate the gravitational and nongravi-
tational forces on a system of bodies accurately over long
timescales (known as the n-body problem). The solution
to the dynamical evolution of any system of three or more
bodies cannot be solved exactly, but there are many dif-
ferent types of mathematical algorithms that have been
developed to optimize the pursuit of an approximate
solution (i.e., Runge–Kutta, symplectic integration, etc).
Important applications for these tools include the study of
the formation of planetary systems through long-term
accretion of planetary embryos, as well as the dynamical
history of populations of small bodies in our Solar System
(such as the Kuiper Belt).
See also
▶ Apsidal Angle
▶ Gravitation
▶ Kuiper Belt
▶ Orbit
▶ Period
▶ Planet Formation
References and Further Reading
Danby JMA (1992) Fundamentals of celestial mechanics. Willmann-Bell

Orbital Period/Frequency
Definition
The orbital period (P) is the time required for a body to
complete an orbit around a central mass. The orbital
period can be calculated by the Kepler’s third law, which
in the solar system takes the form
P 2 ¼ a3
where P is in years and the orbit’s semimajor axis a is in
AU. For different stellar masses the full equation that
should be used is
4p2 a3
P2 ¼
Kirkwood announced the discovery of gaps in the
G Mstar þ Mp
where Mstar and Mp are the stellar and planetary mass,
respectively, and G is the gravitational constant.
See also
▶ Mean Motion Resonance
▶ Orbit
▶ Secular Dynamics
Orbital Resonance
JÉRÔME PEREZ
Applied Mathematics Laboratory, ENSTA ParisTech, Paris
Cedex 15, France
Keywords
Gravitation, instability, period, orbits
Definition
An orbital resonance is either a stabilizing or disrupting
phenomenon produced by the cumulative effect of gravity
between at least two bodies in periodic orbits, typically
around a third, more massive body.
Overview
The motion of a body in a fixed gravitational field is
described by an ordinary differential equation. The solu-
tion of this problem is entirely determined provided we
know its initial conditions.
In several physical cases, this solution has some peri-
odic characteristics. The period of such a periodic motion
depends essentially on a mean distance between bodies in
Organelle O 1181
gravitational interaction. For the two-body Keplerian
problem, this is Kepler’s third law: The square of the
orbital period is directly proportional to the cube of the
semi-major axis of its elliptical orbit (see ▶ Gravitation).
If several bodies are in orbit in a fixed gravitational
potential and the ratio of the period of two of these
particular motions is rational, a special configuration
occurs periodically. The cumulative effects of this config-
uration could be favorable for reaching a stabilized state
or, on the contrary, could lead to a destructive process of
the system or of part of it, by ejection.
The solar system contains plenty of orbital resonances.
Between Mars and Jupiter, there exist thousands
of small bodies that form the asteroid belt. In 1866,
distances of these bodies’ orbits from the Sun. These
gaps were located at positions where, if there were
a body, its period of revolution around the Sun would be
an integer fraction of Jupiter’s orbital period. In fact, these
gaps were dug by orbital resonances. Likewise, Mimas,
a minor satellite of Saturn, has a period that is twice the
one of a body, which would be found in the Cassini
division of the planet’s ring.
Sometimes resonances can maintain rather than
exclude bodies from their orbits. Io, Europa, and Gany-
mede, the first three moons of Jupiter, show a special kind
of orbital resonance called Laplace’s resonance: their
orbital period are in the ratio 1:2:4. In consequence, these
three moons cannot be aligned together on the same side of O
Jupiter. This configuration could eject one of them.
See also
▶ Gravitation
▶ Lagrangian Points
References and Further Reading
Carl DM, Stanley FD (1999) Solar system dynamics. Cambridge
University Press, Cambridge
Organelle
Definition
An organelle is a specialized subunit inside a cell. It is
separated from the surrounding cytoplasmic media by
a lipid bilayer and plays a particular role or function
within the cell similar to that of organs in an animal’s
body. The absence of a ▶ nucleus, a membrane-enclosed
organelle containing the genome, is one of the major
1182 O Organic Cyanide
features that differentiate prokaryotes from eukaryotes.
The most notable examples of organelles are those
involved in energy transduction reactions in eukaryotes
which were originated by endosymbiont bacteria: ▶ chlo-
roplasts present in plants cells, algae and some protists
which are responsible for ▶ photosynthesis, and ▶ mito-
chondria in almost all eukaryotes where ▶ respiration for
energy production takes place. Other important organ-
elles are hydrogenosomes involved in energy generation in
anaerobic conditions; the endoplasmic reticulum respon-
sible for many functions, including the synthesis of new
membrane material; lysosomes involved in protein degra-
dation and amino acid recycling; peroxisomes responsible
for the degradation of oxidative stress products, like
hydrogen peroxide; microfilaments and microtubules
responsible for the internal structure of eukaryotic cells;
and flagella and cilia involved in cell movement.
See also
▶ ATP Synthase
▶ Chloroplast
▶ Eukarya
▶ Mitochondrion
▶ Motility
▶ Nucleus
▶ Oxygenic Photosynthesis
▶ Photosynthesis
▶ Respiration
Organic Cyanide
▶ Nitrile
Organic Material Inventory
Definition
The organic material inventory is the list of materials that
contain organic compounds (typically carbon, hydrogen,
oxygen nitrogen. . .) that are carried on spacecraft in quan-
tities above a specified limit. The list comprises the denom-
ination of each compound and the total amount present
inside the spacecraft. This inventory is a component of
required ▶ planetary protection documentation for mis-
sions to target objects for which there is an interest in
studying organic compounds that could have contributed
to the origin and evolution of life in the Solar System.
Organic Molecule
HENDERSON JAMES (JIM) CLEAVES II
Geophysical Laboratory, Carnegie Institution of
Washington, Washington, DC, USA
Definition
An organic molecule is any member of a large class of
molecules containing carbon, and then limited by a num-
ber of somewhat arbitrary restrictions. For historical rea-
sons, a strict definition of an organic molecule is difficult.
The word “organic” dates back to the ancient Greeks. For
centuries, many in the West believed in the concept of
vitalism: that certain “organic” compounds could only be
synthesized by the action of a vital “life-force” possessed
only by living organisms. This implied that “organic”
compounds were fundamentally different from the “inor-
ganic” compounds that could be obtained by laboratory
manipulation.
Overview
The first well-documented experimental synthesis of an
“organic compound” occurred in 1828 when Wöhler syn-
thesized ▶ urea from the “inorganic” compounds potas-
sium cyanate and ammonium sulfate. Urea had been
considered an “organic” compound, as it was only
known to occur in the urine of living organisms. Since
then, many thousands of biological molecules have been
synthesized by organic chemists.
Some sources define organic compounds as those
containing C-H bonds; others include C-C bonds in the
definition, still others merely require a molecule to
contain carbon. This last definition would include com-
pounds such as steel alloys and calcite, which most organic
chemists would not consider part of their field of study.
The C-H bond or C-C bond containing definitions would
exclude urea, and many other compounds commonly
accepted by chemists as “organic.”
While the definition of what constitutes an organic
compound can be ambiguous for a few small molecules
such as urea, CO, and HCN, this is much less true for even
slightly larger molecules. Indeed, as of 2010, many mil-
lions of “organic” compounds have been synthesized by
chemists, and the theoretical number possible is
infinite. For example, the number of small organic
molecules in what is considered the pharmacologically
relevant size range (molecular weight  <500 amu) is
estimated by some to be on the order of 1060.
 Origin of Life O 1183

See also
▶ Carbon Organometallic
▶ Urea
Definition
Organometallic compounds contain carbon atoms
bonded to a metal, often covalently. Many organometallic
complexes feature coordination bonds between a metal
Organic Refractory Matter and an organic ligand. The organic ligand often binds
the metal through a heteroatom such as oxygen or nitro-
Synonyms gen, in which case such compounds are considered coor-
ORM dination compounds. However, if any of the ligands form
a direct metal–carbon bond, then the complex is usually
Definition considered organometallic. There are many naturally
ORM is complex, nonvolatile organic matter. In meteor- occurring organic coordination compounds. For example,
ites, such material is normally called insoluble organic the ▶ protein hemoglobin contains four ▶ heme groups
matter (IOM) and is broadly defined to have low to no in which an iron ion is coordinated to the nitrogen atoms
solubility in a weak acid bath and is also referred to as of the porphyrin rings. Similarly, magnesium is coordi-
protokerogen or humin. Many models of ▶ interstellar nated via nitrogen atoms at the center of ▶ chlorophyll.
dust include an organic refractory mantle over a silicate In contrast, the ▶ coenzyme methylcobalamin, which has
core, with the ORM produced by ultraviolet or cosmic ray a cobalt–methyl bond, is a true organometallic complex.
irradiation of an accreted icy grain mantle. Organic refrac-
tory matter can be synthesized in the laboratory by See also
bombarding ices containing species thought to be present ▶ Chlorophylls
in comets and ▶ interstellar ices (i.e., water, methane and ▶ Coenzyme
ammonia) with high-energy photons or particles. ▶ Heme
▶ Protein
See also
▶ Interstellar Dust
▶ Interstellar Ices O
▶ Tholins
Origin of Life
ANTONIO LAZCANO
Facultad de Ciencias, Universidad Nacional Autónoma de
México (UNAM), Cd. Universitaria, Mexico D.F, Mexico
Organic Residue
▶ Tholins Synonyms
Abiogenesis; Biopoesis; Building blocks of primitive life;
Early life; Emergence of life

Organicism Keywords
Emergence and complexity, heterotrophic hypothesis,
▶ Vitalism Oparin–Haldane, origin of life, prebiotic evolution, pri-
mordial autotrophy, spontaneous generation

Definition
In evolutionary biology, the phrase “origin of life” refers to
Organized Element the first appearance of living entities. If the emergence of
the biosphere is seen as the evolutionary transition
▶ Globule (Nanoglobule) between the nonliving and the living, then it may be
1184 O Origin of Life
meaningless to attempt to draw a strict line between these
two worlds, and the appearance of life on Earth may
therefore be better envisioned as a continuum that
seamlessly joins the prebiotic synthesis and accumulation
of organic molecules in the primitive environment with
the emergence of self-sustaining, replicative chemical sys-
tems capable of undergoing Darwinian evolution.
Overview
The term “origin of life” has several possible meanings, but
from the perspective of contemporary biology and within
the framework of evolutionary theory it refers to the first
appearance of primordial living entities. It is generally
assumed that since antiquity early philosophers and nat-
uralists appealed to spontaneous generation to explain the
origin of life. However, with few exceptions, this was not
the case. Spontaneous generation was seen mostly as
a nonsexual reproductive mechanism, and it was not
until Erasmus Darwin, Georges Louis Leclerc de Buffon,
and Jean-Baptiste de Lamarck incorporated spontaneous
generation within their transformist views that it was
conceived as the mechanism that had led to the first
appearance of life on our planet (Farley 1977).
Like his predecessors, Charles Darwin surmised that
plants and animals arose naturally from primordial
nonliving matter, but like many others he rejected the
idea that putrefaction of preexisting organic compounds
could lead to the appearance of organisms. Darwin’s let-
ters, notebooks, and few published statements on the issue
demonstrate that although he favored the possibility that
life could appear by natural processes from simple inor-
ganic compounds, his reluctance to discuss it in public was
due at least in part to the recognition that it was difficult to
develop fruitful experimental approaches to address the
problem. Nevertheless, his theory established the frame-
work that would lead to a number of attempts to explain
the origin of life by introducing principles of historical
explanation (Peretó et al. 2009).
Panspermia: Past and Present
Darwin’s Origin of Species was published in 1859, the very
same year in which Pasteur began the experiments that
would lead him to disprove spontaneous generation of
microbes. His results had implications that went well
beyond the limits of academia. Since the times of Lamarck
and Buffon spontaneous generation had been associated
in France not only with evolutionary theory but also with
secular attitudes and radical political views. In such an
entangled atmosphere, spontaneous generation embodied
not only support for evolution and transformist views, but
also implied an anticlerical political stance (Farley 1977;
Fry 2002; Strick 2009).
Pasteur’s results made it difficult to explain the origin
of life by means of spontaneous generation, a conclusion
that was interpreted as a major blow to one of the implicit
assumptions underlying Darwin’s views. Several devoted
materialists like Emil du Bois-Reymond, Karl von Nageli,
and August Weismann continued to support the idea of
spontaneous generation, but others, like Hermann von
Helmholtz, felt that they could sidestep the issue by
assuming that viable microbes had been delivered to the
primitive Earth by meteorites, attempting to keep the
validity of evolution. In fact, toward the end of the nine-
teenth century, the belief that life on Earth had evolved
from extraterrestrial organisms elicited a number of pro-
posed mechanisms that could have transported microbes
between planets (see, e.g., Arrhenius 1908), but little
attention was given to the central issue of their actual
origin (Oparin 1924, 1938; Kamminga 1982; Fry 2002).
The recent revival of the panspermia hypotheses has
not changed this situation. Recent proposals that terres-
trial life arose elsewhere and was transferred to our planet
are based on the well-established exchange of solid mate-
rial among the different bodies in the solar system, the
assumption of more benign environments in some of
these other planets during the early stages of planetary
system, and the surprising resistance of a number of pro-
karyotic species to environmental insults such as high
fluxes of UV light or corpuscular radiation. There are of
course alternative explanations for these observations, but
with formidable disregard for plausibility, the panspermia
hypothesis has been recently restated in a variety of con-
texts. However, its current advocates, like those of the late
nineteenth century, simply transfer the question of the
origin of life to another habitable planet in our galaxy
without providing an explanation of how it happened.
The Origin of Life: A Question Without
an Answer?
A century and a half after Darwin admitted how little was
understood about the origin of life, we still do not know
when and how the first living beings appeared on Earth.
Despite the seemingly insurmountable obstacles sur-
rounding the understanding of the origin of life, or per-
haps because of them, there has been no shortage of
discussion about how it took place. Since the attributes
of the first living entities are unknown, it is not surprising
that an inventory of current views on the origin of life
reveals a mixture of opposites of every kind.
Looking back across time since the origin of life
occurred is a scientific exercise fraught with guesswork,

since the evidence required to describe the prebiotic envi-
ronment and the nature of the events that led to the first
living systems is scant and difficult to understand. Since
most of the rocks from early Archean times that have been
preserved have been metamorphosed to a considerable
extent, there is no direct evidence of the environmental
conditions on the Earth at the time of the emergence of
life, nor any fossil register of the predecessors of the first
cells. There is no direct information of the chemical com-
position of the terrestrial atmosphere during the period of
the origin of life, nor of other general and local environ-
mental conditions, which may have been important for
the appearance of living systems (Bada and Lazcano 2009).
Moreover, any explanation of the origin of living sys-
tems should attempt, at least implicitly, the definition of
a set of minimal criteria for what constitutes a living
organism. However, this has proven to be an elusive intel-
lectual endeavor, though not for lack of trying. The
absence of such a definition sometimes gives the impres-
sion that what is meant by the origin of life is described in
somewhat imprecise terms, and that several entirely dif-
ferent and even opposing questions are often confused
(Lazcano 2008). Prior to the discovery of catalytic activity
of RNA molecules, for instance, the origin of the genetic
code and of protein synthesis was considered synonymous
with the appearance of life itself. This is no longer
a dominant point of view: four of the central reactions
involved in protein biosynthesis are catalyzed by
ribozymes, and their complementary nature suggests
that they first appeared in an ▶ RNA world (Yarus 2010),
that is, that ribosome-catalyzed, nucleic acid-coded pro-
tein synthesis is the outcome of Darwinian selection of
RNA-based biological systems, and not of mere physico-
chemical interactions that took place in the prebiotic
environment.
Phylogenomics and Extant Metabolism Do
Not Explain the Origin of Life
The variations of traits common to extant species can be
explained as the outcome of divergent processes from
ancestral life forms that existed prior to the separation of
the three major biological domains, that is, the last com-
mon ancestor of Bacteria, Archaea, and Eucarya. This
entity was likely far removed, if not in geological time, at
least in complexity from the first living systems. Although
no evolutionarily intermediate stages or ancient simplified
versions of the basic biological processes have been dis-
covered in contemporary organisms, the differences in the
structure and mechanisms of gene expression and replica-
tion among Bacteria, Archaea, and Eucarya have provided
insights into the stepwise evolution of the replication and
Origin of Life O 1185
translation apparatus, including some late steps in the
development of the ▶ genetic code. Such comparative
approaches have made it possible to distinguish the ori-
gin-of-life problem from a whole series of other issues,
often confused, that belong to the domain of the early
evolution of microbial life.
Molecular phylogenies, comparative genomics, and
bioinformatics approaches fail to provide direct informa-
tion on the nature of the first living beings. A cladistic
approach to the origin of life is not feasible, since all
possible intermediates that may have once existed have
long since vanished. Phylogenetic analyses based on com-
parative genomics provide important clues to early stages
of biological evolution, but at the time being their appli-
cability cannot be extended beyond a threshold that cor-
responds to a period of cellular evolution in which protein
biosynthesis was already in operation, that is, the RNA/
protein world (Becerra et al. 2007).
Although there have been attempts to deduce the
nature of the first living systems from extant metabolism
(Wächtershäuser 1988), there is a lack of simple continuity
between the biosynthetic and the (possible) prebiotic
pathways (Lazcano and Miller 1999). For instance, it is
likely that the Strecker synthesis and the Bucherer–Bergs
reaction played a major role in the prebiotic accumulation
of amino acids, but these abiotic routes are very different
from transamination and the reverse Krebs cycle found in
extant cells. The prebiotic synthesis of purines is from
HCN and its derivatives, and not from glycine, formate, O
and NH3, as in current biology. It is true that the amino
imidazole carboxamide ribotide in the de novo purine
nucleotide biosynthetic pathway is comparable to the
amino imidazole carbonitrile formed in the prebiotic
pathway, and a few other examples are found in the for-
mation of uracil, glutamic acid, pyrroles, pyrimidines, and
acetic acids (Lazcano 2010a). However, the similarities
between these abiotic reactions and their enzyme-
mediated counterparts in biosynthetic pathways do not
necessarily indicate an evolutionary continuity between
prebiotic chemistry and biochemical pathways, but may
simply reflect chemical determinism. In other words, the
few cases in which some extant metabolic pathways and
prebiotic chemical processes share similar steps may be
due to the unique way in which given reactions can
take place.
From the Primitive Soup to the RNA World
It is generally believed that after Louis Pasteur had
disproved the spontaneous generation of microbes using
his famous swan-necked flask experiments, the discussion
of life’s beginning had been banished to the realm of
1186 O Origin of Life
useless speculation. However, scientific literature from the
first part of the twentieth century shows the many
attempts by major scientists to solve this problem. The
list covers a rather wide range of explanations that go from
the ideas of Pflüger on the role of hydrogen cyanide on the
origin of life, to those of Svante Arrhenius on panspermia,
and includes Leonard Troland’s hypothesis of a primordial
enzyme formed by chance events in the primitive ocean,
Alfonso L. Herrera’s sulfocyanic theory on the origin of
cells, Harvey’s 1924 suggestion of an heterotrophic origin
in a high-temperature environment, and the provocative
1926 paper that Hermann J. Muller wrote on the abrupt,
random formation of a single, mutable gene endowed
with catalytic and autoreplicative properties (cf. Lazcano
1992; 2010b).
In spite of their diversity, most of these explanations
went unnoticed, in part because they were incomplete,
speculative schemes largely devoid of direct evidence and
not subject to fruitful experimental testing. Although
some of these hypotheses considered life as an emergent
feature of nature and attempted to understand its origin
by introducing principles of historical explanation, the
dominant view was that the first forms of life had been
photosynthetic microbes endowed with the ability to fix
atmospheric CO2 and to use it with water to synthesize
organic compounds. A major scientific breakthrough
occurred, however, when Oparin (1924) suggested
a heterotrophic origin of life that assumed that prior to
the emergence of the first cells a prebiotic synthesis of
organic compounds led to the accumulation of the prim-
itive broth.
Such ideas were supported not only by the evidence of
organic compounds in meteorites, but also by the striking
nineteenth-century experimental demonstrations that
biochemical compounds such as urea, alanine, and sugars
could be formed under laboratory conditions, as had been
demonstrated by Wöhler, Strecker, and Butlerow, respec-
tively. Oparin’s proposal, which was based on his Darwin-
ian credence in a gradual, slow evolution from the simple
to the complex, stood in sharp contrast with the then
prevalent idea of an autotrophic origin of life. Since
a heterotrophic anaerobe is metabolically simpler than
an autotrophic one, the former would necessarily have
evolved first. Thus, based on the simplicity and ubiquity
of fermentative reactions, Oparin (1924) suggested in
a small booklet that the first organisms must have been
heterotrophic bacteria that could not make their own food
but rather consumed organic material present in the prim-
itive milieu.
These ideas were further elaborated and refined in
a more extensive book whose English translation was
published in 1938 (Oparin 1938). In this book, Oparin’s
original proposal was revised, leading to the assumption of a
highly reducing milieu in which iron carbides of geological
origin would react with steam to form hydrocarbons. Their
oxidation would yield alcohols, ketones, aldehydes, etc., that
would then react with ammonia to form amines, amides,
and ammonium salts. The resulting protein-like com-
pounds and other molecules would form a hot dilute
soup, in which they would aggregate to form colloidal
systems such as coacervates, from which the first heterotro-
phic microbes evolved. Like many others at the time, Oparin
did not address in his 1938 book the origin of nucleic acids,
because their role in genetic processes was not even
suspected. Because of this, inheritance of primordial genetic
information was assumed by Oparin to be the result of
growth and division in the coacervate drops he advocated
as models of precellular systems (Lazcano 2010b).
Similar ideas were being developed independently by
other researchers. In 1929, John B. S. Haldane, the extraor-
dinary British polymath, also argued that the origin of life
had been preceded by the synthesis of organic compounds.
Based on experiments by E. C. C. Baly, an English chemist
who claimed that he achieved the syntheses of amino acids
and sugars by the UV irradiation of a solution of CO2 in
water, Haldane suggested that the absence of oxygen in
a CO2-rich primitive atmosphere had led to the synthesis
of organic compounds and the formation of a “hot dilute
soup.” Haldane was also influenced by D’Herelle’s discov-
ery of phages, and suggested that viruses represented an
intermediate step in the transition from the prebiotic soup
to the first heterotrophic cells. Life may have remained,
wrote Haldane (1929), in the viral stage for many millions
of years before leading to the first cells.
The hypothesis that the first organisms were anaerobic
heterotrophs is based on the assumption that abiotic
organic compounds were a necessary precursor for the
appearance of life. Experimental evidence in support of
Oparin’s proposal of chemical evolution came first from
Harold C. Urey’s laboratory, whom had been involved
with the study of the origin of the solar system and the
chemical events associated with this process. Urey had also
considered the origin of life in the context of his proposal
of a highly reducing terrestrial atmosphere (Urey 1952).
The first successful prebiotic amino acid synthesis was
carried out with an electric discharge and a strongly reduc-
ing model atmosphere of CH4, NH3, H2O, and H2 (Miller
1953). The result of this experiment was a large yield of
a racemic mixture of amino acids, together with hydroxy
acids, short aliphatic acids, and urea. One of the surprising
results of this experiment was that the products were not
a random mixture of organic compounds; rather,

a relatively small number of compounds were produced in
substantial yield. Moreover, with a few exceptions, the
compounds were of biochemical significance.
It is very unlikely, however, that the RNA world would
have arisen from such abiotic organic syntheses. The dis-
covery of catalytically active RNA molecules gave consid-
erable credibility to prior suggestions that the first living
organisms were largely based on ribozymes, a hypothetical
stage called the RNA world (Gilbert 1986; Joyce 2002).
This possibility is now widely accepted, but the problems
involved with the synthesis and accumulation of RNA
components suggest that ribozymes were not a direct out-
come of prebiotic evolution, but may have been one of the
evolutionary outcomes of what are now referred to as pre-
RNA worlds. However, the chemical nature of the first
genetic polymers and the catalytic agents that may have
formed the pre-RNA worlds that bridged the gap between
the prebiotic broth and the RNA world are completely
unknown and can only be surmised.
The catalytic versatility of RNA molecules clearly
merits a critical reappraisal of Muller’s viewpoints (Muller
1961). However, the discovery of ribozymes does not
imply that autocatalytic nucleic acid molecules ready to
be used as primordial genes were floating in the primitive
oceans, or that the RNA world emerged completely assem-
bled from simple precursors present in the prebiotic soup.
The evidence supporting the presence of a wide range of
organic molecules on the primitive Earth, including mem-
brane-forming compounds, suggests that the evolution of
membrane-bounded molecular systems preceded cellular
life on our planet, and that life is the evolutionary out-
come of a process, not of a single, fortuitous event.
A Modern Scientific Battlefield
As summarized by Eschenmoser (2008), two major camps
can be recognized among those working on the origins of
life, that is, those assuming that the emergence of auto-
catalytic “metabolic” cycles in the primitive Earth was
essential for the appearance of genetic systems, and those
that assume the priority of genetic polymers endowed
with catalytic properties. These two different viewpoints
reflect a rather sharp division between those who favor the
idea that life is an emergent interactive system endowed
with dynamic properties that exist in a state close to
chaotic behavior, and those who are reluctant to adhere
to a definition of living systems lacking of a genetic com-
ponent whose properties reflect the role that Darwinian
natural selection and, in general, evolutionary processes,
have played in shaping its central characteristics.
With few exceptions, such as the views advocated by
Sidney W. Fox and others (cf. Fox and Dose 1977), during
Origin of Life O 1187
the years that followed the ▶ Miller–Urey experiment
attempts to understand the origin of life were shaped to
a considerable extent by the unraveling of the molecular
details of DNA replication and protein biosynthesis.
Perhaps not surprisingly, Herman J. Muller profited
from the rapid developments in molecular biology and
the success in prebiotic syntheses to update his gene-first
proposal by arguing that what had emerged in the prim-
itive oceans had been, in fact, a primordial DNA. For
Muller and his followers, the essence of life lies in the
combination of autocatalysis, heterocatalysis, and muta-
bility, which he equated with evolutionary potential. In
doing so, he implicitly stated that life could be so well
defined that the exact point at which it started could be
established with the sudden appearance of the first DNA
molecule (Muller 1961).
On the other hand, Wächtershäuser (1988) has argued
that life started with the appearance of an autochemo-
lithotrophic two-dimensional monomolecular layer bound
to pyrite that lacked a genetic system. According to this
proposal, the synthesis in activated form of organic com-
pounds such as amino acid derivatives, thioesters, and
keto acids is assumed to have taken place on the surface
of FeS and FeS2 in environments that resemble those of
deep-sea hydrothermal vents. Wächtershäuser’s explicit
adherence to Karl Popper’s philosophical stance (Popper
1959) played a major role in shaping his ideas. The
hypothetico-deductive system developed by Popper under-
lines at least in part Wächtershäuser’s so-called methodol- O
ogy of retrodiction, which he has used to argue that “lines
of descent can be traced back to the first archaic pathways.”
According to this scheme, chemoautotrophic carbon fixa-
tion took place by a reductive citric acid cycle, or reverse
Krebs cycle, of the type originally described for the photo-
synthetic green sulfur bacterium Chlorobium limicola.
Wächtershäuser’s insightful prediction that ferrous
sulfide in the presence of hydrogen sulfide (H2S) is an
efficient reducing agent should not be underappreciated.
Pyrite formation can produce molecular hydrogen, pro-
mote the formation of ammonia from dinitrogen, and can
reduce a few organic molecules under relatively mild
conditions. However, compared with the surprising vari-
ety of biochemical compounds that are readily synthesized
in one-pot Miller–Urey type simulations, the suite of
molecules produced under the conditions suggested by
Wächtershäuser is quite limited. Wächtershäuser’s proposal,
which is part of a trend that assumes that genetic material
was not essential for the origin of life, has been modified by
Russell and Hall (1997) and Martin and Russell (2003),
who have argued that a primitive version of the acetyl-
CoA pathway existed within FeS-rich mineral boundaries.
1188 O Origin of Life
Based on the hypothesis that core metabolic processes
have not changed since the emergence of life, Morowitz
(1992) has also argued that intermediary metabolism
recapitulates prebiotic chemistry. He maintains that the
basic traits of metabolism could only evolve after the
closure of an amphiphilic bilayer membrane into
a vesicle, that is, that the appearance of membranes rep-
resents the discrete transition from nonlife to life.
According to his hypothesis, reverse Krebs cycle-
dependent life appeared with “minimal protocells”
formed by bilayer vesicles made up of small amphiphiles
and endowed with pigments capable of absorbing radiant
energy stored as a chemiosmotic proton gradient across
the membrane.
Life as an Emergent Complex System
The development of metabolic approaches to the origin of
life like that of Morowitz (1992) and his followers is
increasingly based on complexity theories and self-
assembly phenomena. The background of current meta-
bolic views lies not in Oparin’s proposals, but in the
attempt to extrapolate to biology the deeply rooted ten-
dency in physical sciences to search for all-encompassing
laws that can be part of grand theory that can explain
many, if not all, complex systems (Lazcano 2010a). These
explanations of the origin of life are based on the idea
that the emergence of autocatalytic “metabolic” cycles
on the primitive Earth was an essential prerequisite for
the appearance of genetic systems. According to this
approach, life can be considered an emergent interactive
system endowed with dynamic properties that exist in
a state close to chaotic behavior. Some of these proposals
reflect a reaction against the reductionism of molecular
biology, as well as the adherence to all-encompassing
views based on complexity theories and self-assembly
that tend to be more popular among physical scientists
than among biologists.
Of course, the many examples of self-organizing phys-
ical systems that lead to highly ordered structures demon-
strate that, in addition to natural selection, there are other
mechanisms of ordered complexity that operate. Self-
assembly is not unique to biology, and may indeed be
found in a wide variety of systems, including cellular
automata, the complex flow patterns of many different
fluids, in cyclic chemical phenomena (such as the
Belousov–Zhabotinsky reaction) and, quite significantly,
in the autoorganization of lipidic molecules in bilayers,
micelles, and liposomes. There are indeed some common
features among these systems, and it has been suggested
that “emergent properties” or “self-organizing principles”
suffice to explain the origin of life with the emergence of
self-organizing metabolic cycles that at first did not
require genetic polymers.
The available evidence suggests that the origin of life
and the onset of natural selection resulted from self-
organization phenomena involving nonequilibrium
physical and chemical processes that go beyond simple
Newtonian physics, and that were based on the abiotic
synthesis, stability, and accumulation of organic mole-
cules endowed with differential reactivity, on nucleic
acid self-assembly and chemistry, and the spontaneous
assembly of prebiotic amphiphiles into micelles and
bilayer membranes, among others (de Duve 2005; Lehn
2002; Budin and Szostak 2010). The many examples of
self-organizing physical systems that lead to highly
ordered structures suggest that, in addition to natural
selection, there are other mechanisms of ordered complex-
ity that may have played a role in the origin of living
systems. The recognition that the emergence of life is the
outcome of an evolutionary process constrained by the
laws of physics and chemistry can lead to the acceptance
that many properties associated with living systems, such
as replication, self-assembly, or nonenzymatic catalysis are
also found in nonliving entities (Lazcano 2010a, b). More-
over, if the origin of life is seen as the evolutionary tran-
sition between the nonliving and the living, then it is
meaningless to attempt to draw a strict line between
these two worlds, and the appearance of life on Earth
should, therefore, be seen as an evolutionary continuum
that seamlessly joins the prebiotic synthesis and accumu-
lation of organic molecules in the primitive environment,
with the emergence of self-sustaining, replicative chemical
systems capable of undergoing Darwinian evolution
(Lazcano 2010a, b).
Conclusions
The remarkable coincidence between the monomeric con-
stituents of living organisms and those synthesized in
laboratory simulations of the prebiotic environment
appears to be too striking to be fortuitous. Nevertheless,
at the time being the gap between the primitive soup and
the RNA world is discouragingly large. Life cannot be
reduced to one single molecule such as DNA or a popula-
tion of replicating ribozymes, but present-day biology
indicates that it could not have evolved in the absence of
a genetic replicating mechanism ensuring the stability and
diversification of its basic components.
As shown by the work of Jack Szostak and his associ-
ates (Mansy et al. 2008), it is possible to overcome at least
in part the intellectual dichotomy between those claiming
that the appearance of the first life forms depended
on informational oligomeric compounds, that is, the

so-called genetic approach, and those that argue that it
was based on autocatalytic metabolic cycles. Instead of
engaging in footling arguments about when exactly did
life start, the recognition that it is the outcome of an
evolutionary process constrained by the laws of physics
and chemistry can lead to the acceptance that many prop-
erties associated with living systems, such as replication,
self-assembly, or catalysis are also found in nonliving
entities.
Perhaps we will never know exactly how life origi-
nated. As in other areas of evolutionary biology, answers
to questions on the origin and nature of the first life forms
can only be regarded as inquiring and explanatory rather
than definitive and conclusive. This does not imply that all
origin-of-life theories and explanations can be dismissed
as pure speculation, but rather that the issue should be
addressed conjecturally, in an attempt to construct not
a mere chronology but a coherent historical narrative by
weaving together a large number of observations and
experimental results (Kamminga 1982).
See also
▶ Baly’s Experiment
▶ Bernal’s Conception of Origins of Life
▶ Calvin’s Conception of Origins of Life
▶ Cell
▶ Chemical Evolution
▶ Complexity
▶ Darwin’s Conception of Origins of Life
▶ Evolution (Biological)
▶ Genetics
▶ Life
▶ Metabolism (Biological)
▶ Monod’s Conception on the Origins of Life
▶ Oparin’s Conception of Origins of Life
▶ Panspermia
▶ Phylogeny
▶ Prebiotic Chemistry
▶ Replication (Genetics)
▶ RNA World
▶ Urey’s Conception of Origins of Life
References and Further Reading
Arrhenius S (1908) Worlds in the making: the evolution of the Universe.
Harper & Row, New York
Bada JL, Lazcano A (2009) The origin of life. In: Ruse M, Travis J (eds) The
harvard companion of evolution. Belknap/Harvard University Press,
Cambridge, pp 49–79
Becerra A, Delaye L, Islas A, Lazcano A (2007) Very early stages of
biological evolution related to the nature of the last common ances-
tor of the three major cell domains. Annu Rev Ecol Evol Syst
38:361–379
Origin of Life O 1189
Budin I, Szostak JW (2010) Expanding roles for diverse physical phenom-
ena during the origin of life. Annu Rev Biophys 39:245–263
De Duve C (2005) The onset of selection. Nature 433:581–582
Eschenmoser A (2008) On a hypothetical generational relationship
between HCN and constituents of the reductive citric acid cyle. In:
Herdewijn P, Kisakürek V (eds) Origin of life: chemical approach.
Wiley-VCH, Zürich, pp 31–50
Farley J (1977) the spontaneous generation controversy from descartes to
oparin. John Hopkins University Press, Baltimore/London
Fox SW, Dose K (1977) Molecular evolution and the origin of life. Marcel
Dekker, New York
Fry I (2002) The emergence of life on earth. Rutgers University Press,
New Brunswick
Gilbert W (1986) The RNA world. Nature 319:618
Haldane JBS (1929) The origin of life. Rationalist Annu 148:3–10
Joyce GF (2002) The antiquity of RNA-based evolution. Nature 418:
214–221
Kamminga H (1982) Life from space – a history of panspermia. Vistas
Astron 26:67–86
Lazcano A (1992) Origins of life: the historical development of recent
theories. In: Margulis L, Olendzenski L (eds) Environmental evolu-
tion: effects of the origin and evolution of life on planet earth. MIT
Press, Cambridge, pp 57–69
Lazcano A (2008) What is life? A brief historical overview. Chem Biodivers
5:1–15
Lazcano A (2010a) Which way to life? Orig Life Evol Biosph
40:161–167
Lazcano A (2010b) Historical development of origins of life. In: Deamer
DW, Szostak J (eds) Cold spring harbor perspectives in biology:
the origins of life. Cold Spring Harbor Press, Cold Spring Harbor,
pp 1–16
Lazcano A, Miller SL (1999) On the origin of metabolic pathways. J Mol
Evol 49:424–431
Lehn JM (2002) Towards self-organization and complex matter. Science
295:2400–2403
O
Mansy SS, Schrum JP, Krishnamurthy M, Tobé S, Treco DA, Szostak JW
(2008) Template-directed synthesis of a genetic polymer in a model
protocell. Nature 454:122–125
Martin W, Russell MJ (2003) On the origin of cells: a hypothesis for the
evolutionary transition s from abiotic chemistry to chemoautotro-
phic prokaryotes, and from prokaryotes to nucleated cells. Philos
Trans R Soc Lond B Biol Sci 358:59–85
Miller SL (1953) A production of amino acids under possible primitive
Earth conditions. Science 117:528
Morowitz HJ (1992) Beginnings of cellular life: metabolism recapitulates
biogenesis. Yale University Press, New Haven
Muller HJ (1961) Genetic nucleic acid: key material in the origin of life.
Persp Biol Med 5:1–23
Oparin AI (1924) Proiskhozhedenie Zhizni. Mosckovskii Rabochii,
Moscow. Reprinted and translated in Bernal JD (1967) The origin
of life. Weidenfeld and Nicolson, London
Oparin AI (1938) The origin of life. McMillan, New York
Peretó J, Bada JL, Lazcano A (2009) Charles Darwin and the origins of life.
Orig Life Evol Biospheres 39:395–406
Popper K (1959) The logic of scientific discovery. Basic Books,
New York
Russell MJ, Hall AJ (1997) The emergence of life from iron monosulphide
bubbles at a submarine hydrothermal redox and pH front. J Geol Soc
Lond 154:377–402
Strick JE (2009) Darwin and the origin of life: public versus private
science. Endeavour 33(4):148–151
1190 O ORM

Urey HC (1952) The planets: their origin and development. University of See also
Chicago Press, Chicago
▶ Amino Acid
Wächtershäuser G (1988) Before enzymes and templates: theory of surface
metabolism. Microbiol Rev 52:452–484
▶ Arginine
Yarus M (2010) Life from an RNA World: the ancestor within. Harvard ▶ Hydrolysis
University Press, Cambridge ▶ Peptide

References and Further Reading

Meierhenrich UJ, Muñoz Caro GM, Bredehöft JH, Jessberger EK,
Thiemann WH (2004) Identification of diamino acids in the Mur-
ORM chison meteorite. Proc Natl Acad Sci USA 101:9182–9186. Available
at: http://www.pnas.org/cgi/doi/10.1073/pnas.0403043101

▶ Organic Refractory Matter

Orpheus
Ornithine ▶ Theia

Synonyms
Diaminovaleric acid
Definition
L-Ornithine is a natural, biological, non-coded diamino
acid (with a and d-amino groups). L-Ornithine (Fig. 1)
was first isolated by Jaffé from chicken excrement in 1877,
hence its name (from Greek: orniς (ornis) = bird). It plays
a major role as a carrier compound in the metabolic urea
cycle. Ornithine can be derived from glutamate and is used
in the biosynthesis of arginine and is therefore indirectly
coupled to protein synthesis in mammals. Polypeptides
containing unprotected ornithines undergo spontaneous
lactamization. L-ornithine is produced through hydrolysis
of citrulline and L-arginine in alkaline medium, that is, the
reverse of the biosynthetic process or the same direction
observed in the urea cycle. Both, L- and D-ornithine have
been tentatively identified in the Murchison meteorites
(Meierhenrich 2004), (concentration < 5ppb).
Molecular formula: C5H12N2O2
Molecular mass: 132.2 g·mol1
O
H2N OH
NH2
Ornithine
Ornithine. Figure 1
Ortholog
▶ Orthologous Gene
Orthologous Gene
Synonyms
Ortholog
Definition
Orthologous genes (or orthologs) are a particular class of
homologous ▶ genes. They are found in different species
and have diverged following the speciation of the species
hosting them. Therefore, orthologous genes in different
species derive from a ▶ common ancestral gene found in
the ancestor of those species. Given their common origin,
it is often the case that orthologous genes have the same
function in the different species, but exceptions are not
rare. The best way to infer that genes from different species
are orthologous is by reconstructing their evolutionary
relationships using molecular ▶ phylogeny. Conversely,
because orthologous genes evolve in parallel with the
diversification of species, they are markers of choice for
the reconstruction of the evolutionary history of species
using molecular phylogeny.
See also
▶ Common Ancestor
▶ Gene
 OTU O 1191

▶ Homology
▶ Paralogous Gene Osmotic Pressure
▶ Phylogeny
Definition
▶ Water diffuses through semipermeable ▶ membranes
from low- to high-solute- concentration solutions. This
Orthophosphate process is known as osmosis. Osmotic pressure is the
pressure produced on the semipermeable membrane of
▶ Phosphoric Acid an organism by the difference in solute concentration
between the internal and the external part of the cell.
When the solute concentration in the interior of the cell
is as that of the medium, we say that the solution is
isotonic, no diffusion takes place. Normally, cell systems
Osmolite develop in environments where the solute concentration is
lower than the cytoplasmic solute concentration. In this
Synonyms case, water diffuses from the medium to the cytoplasm.
Osmolyte
Halophilic organisms grow in solutions where the solute
concentration is much higher than the cytoplasmic one. In
Definition this case, there is a tendency for water to flow out of the
Osmolites or ▶ compatible solutes are organic molecules
cell to diminish the osmotic pressure on the membrane.
which are compatible with cell metabolism even at molar
This can produce severe problems for cells if they have no
concentrations and play protective roles in adaptation to
means with which to deal with it. Normally, halophilic
extreme environments. Microorganisms also accumulate
organisms have cytoplasmic ▶ compatible solutes,
organic solutes for osmotic adjustment. ▶ Thermophilic
organic or inorganic, which try to compensate the high
and ▶ hyperthermophilic organisms accumulate alcohol
osmotic pressure generated by the ionic strength of the
phosphates (e.g., inositol-phosphates) as compatible sol-
medium in which to develop.
utes for thermal stabilization, something which does not
occur in bacteria or archaea that grow at low or moderate
temperature. Osmolites or soluble compatibles can be See also O
classified into three groups: poly-alcohol molecules (e.g., ▶ Archea
manytol), quaternary ammines (e.g., glycine betaine), and ▶ Bacteria
amino acids (e.g., proline). Compatible solutes have been ▶ Compatible Solute
studied in vitro and have shown significant effects on ▶ Extremophiles
biomolecules as stabilizers of native macromolecules (pro- ▶ Eukarya
teins or nucleic acid structures). ▶ Halophile
▶ Halotolerance
See also ▶ Membrane
▶ Compatible solute ▶ Osmolite
▶ Enzyme
▶ Extremophiles
▶ Halophile
▶ Halotolerance
▶ Hyperthermophile OST
▶ Thermophile
▶ Outer Space Treaty

Osmolyte
OTU
▶ Compatible Solute
▶ Osmolite ▶ Phylotype
1192 O Outer Membrane
Outer Membrane
Definition
The outer ▶ membrane is a phospholipid- and polysac-
charide-containing membrane that lies external to the
▶ peptidoglycan layer of ▶ gram-negative bacteria. This
layer is effectively a second lipid bilayer, but in addition to
phospholipids and proteins found in the cytoplasmic
membrane, it contains polysaccharides. It is a rather com-
plex structure only found in gram-negative bacteria.
The lipid and polysaccharides are linked to form
a lipopolysaccharide complex. For this reason, the outer
membrane is often called the lipopolysaccharide layer
(LPS). In the outer membrane, LPS associates with several
proteins to form the outer leaflet of the membrane. Lipo-
proteins are also found in the inner leaflet of the outer
membrane and their function is to anchor the membrane
to the peptidoglycan. In the outer part of the membrane
lipopolysaccharides replace phospholipids. Thus, although
the outer membrane can be considered a lipid bilayer, its
structure and function is quite distinct from that of the
cytoplasmic membrane. Although the major function of
the outer membrane of gram-negative bacteria is struc-
tural, its components can produce toxicity to animals. The
outer membrane has special types of proteins called
porins, which are channels responsible for the permeabil-
ity of small molecules. The periplasm is the space
delimited by the outer layer and the cytoplasmic mem-
brane of the gram-negative bacteria in which different
important cellular functions are performed.
See also
▶ Gram Negative Bacteria
▶ Membrane
▶ Peptidoglycan
Outer Space Treaty
Synonyms
OST
Definition
The “Treaty on Principles Governing the Activities of
States in the Exploration and Use of Outer Space, includ-
ing the Moon and Other Celestial Bodies” represents the
basic legal framework of international space law. It was
proposed to the UN by the USSR and the US in 1966, and
ratified by those countries and, to date, up to 96 others
starting in 1967. The Outer Space Treaty (OST) sets up the
general principles applicable to the exploration and use of
outer space and applies to every activity of states in outer
space, whether such activities are carried out by govern-
mental or nongovernmental entities. The Article IX, sec-
ond sentence, states: “State Parties shall pursue studies of
outer space, including the Moon and other celestial bod-
ies, and conduct exploration of them so as to avoid their
harmful contamination and also adverse changes in the
environment of the Earth resulting from the introduction
of extraterrestrial matter and, when necessary, adopt
appropriate measures for this purpose.” The ▶ COSPAR
refers to this article as the legal basis for the ▶ planetary
protection policy on which all subsequent recommenda-
tions and guidelines are based.
See also
▶ COSPAR
▶ Planetary Protection
References and Further Reading
United Nations. Treaty on Principles Governing the Activities of States in
the Exploration and Use of Outer Space, including the Moon and
Other Celestial Bodies (the “Outer Space Treaty”) referenced 610
UNTS 205 – resolution 2222(XXI) of December 1966
Outflow Channels
ERNST HAUBER
German Aerospace Center (DLR), Institute of Planetary
Research, Berlin, Germany
Keywords
Erosion, flooding, groundwater, ice, water
Definition
Outflow channels on ▶ Mars are large channel systems
that emerge fully-sized from rubble-filled “chaotic”
depressions, are generally tens of kilometers across and
hundreds of kilometers long, and were formed by fluid
flow of ▶ water from subsurface reservoirs.
Overview
Outflow channels were first detected on images from the
Mariner 9 mission (Baker and Milton 1974). They are the
strongest morphological evidence that large amounts of
 1,3-Oxazolidine-2,5-dione O 1193

water once flowed on the Martian surface. Characteristic Baker VR, Milton DJ (1974) Erosion by catastrophic floods on Mars and
Earth. Icarus 23:27–41
landforms of outflow channels are anastomosing (build-
Baker VR, Strom RG, Gulick VC, Kargel JS, Komatsu G, Kale VS
ing a network of streams that both branch out and (1991) Ancient oceans, ice sheets and the hydrological cycle on
reconnect) channels, streamlined erosional “islands,” dry Mars. Nature 352:589–594
cataracts, and scour marks and grooves on the channel Bates R, Jackson J (1980) Glossary of geology, 2nd edn. American
floors (Baker 1982). The closest terrestrial analog for out- Geological Institute, 677 pp
Bretz H (1923) The Channeled Scabland of the Columbia Plateau. J Geol
flow channels is the huge complex of anastomosing rock-
31:617–649
cut fluvial channels in east-central Washington (USA). Head JW, Wilson L, Mitchell KL (2003) Generation of recent massive
This so-called “Channeled Scabland” was first interpreted water floods at Cerberus Fossae, Mars by dike emplacement,
by J Harlan Bretz as the result of the catastrophic cryospheric cracking, and confined aquifer groundwater release.
megaflooding derived from ice-dammed Pleistocene Geophys Res Lett 30:1577. doi:10.1029/2003GL017135
Lucchitta BK (1982) Ice sculpture in the Martian outflow channels.
glacial lakes (e.g., Bretz 1923), an interpretation which
J Geophys Res 87:9951–9973
challenged the conventional geologic wisdom of Unifor- Neukum G, Basilevsky AT, Kneissl T, Chapman MG, Van Gasselt S,
mitarianism and was, therefore, heavily debated (Baker Michael G, Jaumann R, Hoffmann H, Lanz JK (2010) The
2009). Uniformitarianism (as described by Bates and Jack- geologic evolution of Mars: episodicity of resurfacing events and
son 1980) is the fundamental principle or doctrine that ages from cratering analysis of image data and correlation with
radiometric ages of Martian meteorites. Earth Planet Sci Lett
geologic processes and natural laws now operating to
294:204–222
modify the Earth’s ▶ crust have acted in the same regular
manner and with essentially the same intensity through-
out geologic time, and that past geologic events can be
explained by phenomena and forces observable today; the
classical concept that “the present is the key to the past.” Outgassing
Many models of the formation of Martian outflow chan-
nels favor a catastrophic formation by the sudden release ▶ Degassing
of large amounts of water from subsurface reservoirs.
Pressurized aquifers might have existed below an imper-
meable frozen upper crust (the ▶ cryosphere), which was
then cracked by the ascent of volcanic dikes (e.g., Head
et al. 2003). Alternatively, outflow channels might have Oxalic Acid Dinitrile O
been eroded, at least partly, by glacial processes (Lucchitta
1982). Most, if not all, outflow channels show evidence for ▶ Cyanogen
a multi-stage formation with different phases of activity
separated by geologically long periods of time. Outflow
channel activity spans a long time, from 3.5 Ga to
geologically very recent times (Neukum et al. 2010). Out-
flow channels terminate into the northern lowlands of
Oxalonitrile
Mars and might have fed an ancient northern ocean
▶ Cyanogen
(Baker et al. 1991), although this scenario is contentious.

See also
▶ Crust
▶ Cryosphere Oxalyl Cyanide
▶ Mars
▶ Valley Networks ▶ Cyanogen
▶ Water

References and Further Reading

Baker VR (1982) The channels of Mars. University of Texas Press, Austin,
198 pp
1,3-Oxazolidine-2,5-dione
Baker VR (2009) The Channeled Scabland: a retrospective. Annu Rev
Earth Planet Sci 37:393–411 ▶ Amino Acid N-Carboxy Anhydride
1194 O Oxic

to copper metal (Cuo). Thus, in the reaction, the reductant

Oxic or ▶ reducing agent loses electrons and is oxidized, and
the oxidant or oxidizing agent gains electrons and is
Definition reduced.
Oxic is a term used to describe a condition, environment,
or habitat in which oxygen is present. Oxidation in Terms of Hydrogen Transfer
Oxidation is the loss of hydrogen.
● Methanol (alcohol) can be oxidized to methanal
(aldehyde) by loss of hydrogen
Oxidant
CH3 OH ! H2 CHO
▶ Electron Acceptor
Oxidation in Terms of Oxygen Transfer
Oxidation is the gain of oxygen.
● The reaction between elemental iron and oxygen to
Oxidation form iron oxide involves the oxidation of iron (com-
monly known as rusting)
CONCEPCIÓN ALONSO 4 Fe þ 3 O2 ! 2 Fe2 O3
Universidad Autonoma de Madrid, Madrid, Spain
In general terms, oxidation implies an increase in an atom’s
oxidation state. However, it is only correct to speak about
Synonyms the oxidation state of a metal atom when the compound is
Combustion; Corrosion; Rusting purely ionic. Molecules, atoms or ions that have the ability
to reduce others are said to be reductive and are known as
Keywords reducing agents, reductants, or reducers. That is to say, the
Electron donor, reducing agent, reductive reductant transfers electrons to another substance, which
is, as a result, oxidized. Since, it “donates” electrons it is
Definition also called an ▶ electron donor. The electronic transfer
Oxidation can be defined with different terms such as: the reactions known as redox reactions are especially impor-
loss of electrons or hydrogen, or the gain of oxygen by an tant in biology because most biochemical reactions imply
atom or a molecule. reduction or oxidation reactions.
Substances that act as reductants include, among
Overview others, electropositive elemental metals like lithium,
sodium, magnesium, iron, zinc and aluminum. These
Oxidation in Terms of Electron Transfer metals readily donate or give away electrons. Hydride
Oxidation is the loss of electrons. transfer reagents, such as NaBH4 and LiAlH4, are widely
For example, in the reaction between iron and copper used in organic chemistry. Another method of reduction
sulfate solution involves the use of hydrogen gas (H2) with a palladium,
platinum, or nickel catalyst. These catalytic reductions are
Fe þ CuSO4 ! Cu þ FeSO4 primarily used in the reduction of carbon-carbon double
copper sulfate and iron sulfate are both ionic compounds. or triple bonds.
If rewritten as an ionic equation, it turns out that sulfate
ions are spectator ions and the reaction can be written as: See also
Fe þ Cu 2þ
! Cu þ Fe 2þ ▶ Electron Donor
▶ Redox Potential
where the iron is oxidized from metal iron (Feo) to ferrous ▶ Reducing Agent
iron (Fe2+) by the loss of two electrons:
Fe ! Fe2þ þ 2e References and Further Reading
Encyclopedia Britannica (1992) Macropedia vol 15, pp 1059. Micropedia
while the copper (II) ion, gaining two electrons is reduced vol 9, pp 31, 15th edn. ISBN 0-85229-553-7

Gold V (1997) In: McNaught AD, Wilkinson A (eds) The compendium
of chemical terminology published by the international union
of pure and applied chemistry (IUPAC), 2nd edn. ISBN 0-86542-
684-8
http://chemed.chem.purdue.edu/genchem/topicreview/bp/ch9/redox.php
http://www.chemguide.co.uk/inorganic/redox/definitions.html#top
Hudlický M (1990) Oxidations in organic chemistry. American Chemical
Society, Washington, p 456. ISBN 0-8412-1789-7
Schüring J, Schulz HD, Fischer WR, Böttcher J, Duijnisveld WH (eds)
(1999) Redox: fundamentals, processes and applications. Springer,
Heidelberg, p 246. ISBN 978-3540665281
Oxidative Agent
▶ Electron Acceptor
Oxidative Phosphorylation
▶ Respiration
Oxidizing Atmosphere
Definition
An oxidizing atmosphere is a (planetary) atmosphere
which oxidizes immersed (surface) compounds. It some-
times refers to an O2-rich atmosphere, for example the
atmosphere of modern Earth. Most of the Earth’s atmo-
spheric oxygen is generated by biological photosynthesis;
however, oxidation can also be an effect of photochemistry
initiated by stellar UV radiation. The Martian atmosphere,
for instance, is an oxidizing atmosphere. Its main constit-
uents are CO2, N2, Ar, and H2O but the OH radicals and
oxygen atoms produced by photolysis, result in surface
oxidation and the formation of O2, O3, H2O2.
See also
▶ Oxygenation of the Earth’s Atmosphere
Oxirane
▶ Ethylene Oxide
Oxygen (Atomic) O 1195
Oxygen (Atomic)
Definition
Oxygen is a chemical element with atomic number 8,
represented by the symbol O. The name oxygen was coined
in 1777 by Lavoisier. Its name derives from the Greek word
“oxys” meaning “sharp,” referring to the sharp taste of
acids, and “genēs” meaning “producer.” It is a chalcogen,
and is highly reactive, forming compounds such as oxides
with most other elements. Oxygen is also the name of the
molecular compound O2, also known as ▶ dioxygen,
formed from two covalently linked oxygen atoms with
a spin triplet electron configuration.
Overview
Oxygen is the third most abundant element in the universe
by mass, after hydrogen and helium, and the most abun-
dant element by mass in the Earth’s crust. Dioxygen as
a gas makes up 20.9% of the volume of Earth’s atmo-
sphere, and is also present in the form of water vapor
and CO2. Oxygen makes up 49.2% of the Earth’s crust by
mass, mostly in the form of silicates, carbonates, and metal
oxides, and is the major component of seawater at 88.8%
by mass, in the form of water.
All major classes of biological molecules, including
proteins, nucleic acids, carbohydrates, and fats, contain
oxygen, as do the major inorganic compounds that com-
prise animal shells, teeth, and bone. Oxygen was indepen-
O
dently discovered by Scheele in 1773 or earlier and
Priestley in 1774.
Naturally occurring terrestrial oxygen includes three
stable isotopes: 16O, 17O, and 18O. 16O is the most abun-
dant, making up 99.762% of the natural abundance. Most
16
O is synthesized at the end of the helium fusion process
in stars, but some is made by neon burning. 17O is pri-
marily produced by the burning of hydrogen into helium
during the CNO cycle. 18O is produced when 14N captures
a 4He nucleus during stellar fusion.
Due to its electronegativity, oxygen forms chemical
bonds with almost all other elements to give corresponding
oxides. The ▶ oxidation state of atomic oxygen is 2
in almost all known oxygen-containing compounds, but
in a few compounds such as peroxides the oxidation
state is 1.
See also
▶ Dioxygen
▶ Oxidation
▶ Oxygen Isotopes
1196 O Oxygen (Molecule)

Likewise, the reduction of magnetite, which contains

Oxygen (Molecule) one Fe2+ and two Fe3+, by silica gives fayalite (ferric olivine
Fe2SiO4), which contains two ferrous ions Fe2+:
▶ Dioxygen
2Fe3 O4 þ 3SiO2 , 3Fe2 SiO4 þ O2 ð3Þ
For ideal solids and gas:
PO2 ½Fe2 SiO4 
¼ K2 ðT Þ ð4Þ
Oxygen Catastrophe ½Fe3 O4 ½SiO2 
where concentrations are in bracket. For pure phases,
▶ Great Oxygenation Event
concentrations are equal to 1, while for real gases PO2 is
replaced by fO2. Taking the logarithm of both sides, we get:
DH2
lnfO2 ¼ lnK2 ðT Þ ¼ þ Const ð5Þ
Oxygen Crisis RT
where DH2 is the enthalpy of the reaction and R is the gas
▶ Great Oxygenation Event constant. At a given temperature, the quartz-fayalite-
magnetite (QFM) mineral assemblage (see ▶ Mantle,
Oxidation of entry) imposes or “buffers” a particular
value of fO2. For all practical purpose, oxygen fugacity
measures the electron trade among minerals and describes
Oxygen Fugacity the proportions of Fe2+ and Fe3+ coexisting in the rock,
rather the other way around. Under most conditions, the
FRANCIS ALBARÈDE
fO2 is so low that oxygen is actually fully combined to the
Ecole Normale Supérieure de Lyon, Lyon Cedex 7, France
mineral phases and is not present as a gas. Oxygen fugacity
is a robust estimate of the redox conditions because rocks
are electrical insulators. Other buffers of oxygen fugacity
Keywords exist, such as iron-wüstite (IW), which describes the coex-
Mineral redox buffer; redox reactions; redox zonations
istence of metallic iron Fe0 and Fe2+ in extremely reducing
environments like the lunar mantle.
Definition
Oxygen fugacity is an equivalent of the partial pressure of
See also
oxygen in a particular environment (atmosphere, rocks,
▶ Archean Mantle
etc.) corrected for the nonideal character of the gas.
▶ Earth’s Atmosphere, Origin and Evolution of
▶ Mantle, Oxidation of
Overview ▶ Oxygenation of the Earth’s Atmosphere
There are multiple ways of characterizing how reducing or
▶ Redox Zonation
how oxidizing an environment is. An example of a redox
reaction in an ideal gas mixture of carbon dioxide, meth-
ane, water, and oxygen is:
CO2 þ 2H2 O , O2 þ CH4 ð1Þ
Oxygen Isotopes
The status of this reaction can be evaluated by writing
the equilibrium equation: KO HASHIZUME
PO2 PCH4 Department of Earth and Space Sciences, Osaka
¼ K1 ðT Þ ð2Þ University, Toyonaka, Osaka, Japan
PCO2 PH2 2 O

where Pi stands for the partial pressure of gas i and K1(T )

is the reaction coefficient at temperature T. For real gases, Keywords
pressure Pi should be replaced by fugacities fi. Fugacity and Isotope anomaly, mass-independent-fractionation,
pressure converge toward each other when the total pres- organic matter, photochemistry, self-shielding, solar com-
sure diminishes. position, water

Definition
Oxygen has three stable isotopes, oxygen-16, -17, and -18.
In most studies on terrestrial samples, only the 18O/16O
ratio is usually measured and discussed, where the mass-
dependent fractionation law, expressed by d17O 0.52 
d18O, can be safely assumed. The d17O values among
extraterrestrial samples significantly deviate from the
above shown relationship. This deviation bears the
top-class importance in cosmochemistry, and naturally,
in astrobiology.
Overview
Oxygen is the most abundant element in the planetary
system, except among the giant planets, Jupiter and Sat-
urn, where hydrogen and helium dominate their masses.
Oxygen is accommodated as a major constituent in most
types of building blocks, rocks (MOx), water (H2O), and
organics (CHONS), which are essential in constructing
a terrestrial planet, particularly a habitable planet. Oxygen
isotope compositions among solid planetary materials are
generally expected to bear keys to understand the history
of these building blocks, from their births to their accre-
tion to planets or asteroids, although the details are vig-
orously debated until now (Fig. 1).
Basic Methodology
The oxygen isotope ratios among samples are normally
expressed in the d-notation, that is, by the difference of the
ratios in permil unit relative to the ratios of the reference
standard; dxO
(R/Rstd  1)  1,000; R = xO/16O. The ozone formation in the terrestrial upper atmosphere
international reference standard is the standard mean
ocean water (SMOW) with 17O/16O and 18O/16O ratios
of 3.73  104 and 2.005  103, respectively.
Key Research Findings
Classically, it was considered that rocks and water were
produced from a hot and homogenous solar gas through
equilibrium condensation processes. In this case, we expect
that the O-isotope compositions among the raw material
and all products, that is, the solar and planetary composi-
tions, respectively, are uniformly plotted on a single line
in the d17O versus d18O diagram with a relationship of
d17O 0.52  d18O, namely, the terrestrial fractionation
line (TFL). (Readers are advised to refer to the cross-
references for further basics on the isotope fractionations,
particularly on the mass-dependent fractionations.)
However, in 1973, Clayton and coworkers discovered
that O-isotope compositions among carbonaceous chon-
drites deviate from the TFL, but are plotted around a line
with a much steeper slope of 0.94, so-called the car-
bonaceous chondrite anhydrous minerals (CCAM) line.
Oxygen Isotopes O 1197
Later works (Young and Russell 1998) demonstrated that
the slope is exactly one among the less-altered compo-
nents in the meteorites, but the slope displayed among
assemblages of chondritic minerals actually measured is
slightly smaller, because of the contributions from altered
components, whose O-isotope compositions are some-
what mass-dependently fractionated from their original
compositions. Note that the non-mass-dependent frac-
tionation trend of oxygen is not a peculiar observation
among the carbonaceous chondrites, but are observed
among essentially all available extraterrestrial materials,
including meteorites possibly from Mars or Vesta, one of
the largest asteroids (Clayton 1993, 2005).
Many possible explanations have been submitted to
explain the origin of the slope 1 trend. It was first explained
that the non-mass-dependent O-isotope trend originated
from contributions from a hypothetical 16O-rich exotic
component, possibly a nucleosynthetic origin (Clayton
1993). However, recent measurements of O-isotope com-
positions among pre-solar oxide grains (Nguyen et al.
2007) suggest that the 16O-rich component is rather rare
among the pre-solar grains, thus may not contribute to alter
the bulk meteoritic compositions. The alternative explana-
tions recently supported are the fractionations associated to
chemical processes that took place in the early solar nebula or
in the ancestral molecular cloud. The two different chemical
processes proposed are the mass-independent-fractionation
(MIF) and the CO self-shielding.
MIF is typically seen in the process that operates in the O
(Thiemens 1999). Ozone is formed in the pathway as
follows: O + O2 ↔ O3 ; O3 (+M) ! O3, where O3
denotes the vibrationally excited state, and M is a third
molecule that stabilizes the excited ozone by collision. The
origin of MIF is considered to be the lower density of the
vibrational quantum states for the symmetric isotopomer
(OQO ) compared to the asymmetric one (QOO ), where
Q is either 17O or 18O. The resident time of QOO is longer
than OQO, thus QOO is stabilized more efficiently than
OQO. The QOO excess may occur for both 17O and 18O, by
which the MIF signal is produced. Marcus (2004)
proposed that similar reactions such as O + XO ↔ XO2 ;
XO2 (+M) ! XO2 (X = Si, Ca, Al, etc.), involving MIF,
might have occurred in the early solar nebula, during
the formations of calcium–aluminum-rich inclusions,
which are particularly enriched in 16O, plotted on the
CCAM line.
The CO self-shielding is proposed to take place
during the photodissociation of CO in a gas medium
that consist of H2 and CO, irradiated by external radiation
fields (van Dishoeck and Black 1988; Visser et al. 2009).
1198 O Oxygen Isotopes

600
Slope-1 Line
17,18O-richEndmember in
500 Meteoritic Organic Matter

400
Meteoritic Iron-oxide
(Water Ice Composition)
δ17OSMOW (‰)

300

TFL
Lunar Non-Solar 10
200 Component CI
(Volatile accretion on R
Moon?) 5 L
E
LLH
100 SNC
HED CM
0
CR

−5
CO CV
0 CK
Planetary
CCAM
Proto-Solar −10
−5 0 5 10 15 20
−100
–100 0 100 200 300 400 500 600
δ18OSMOW (‰)

Oxygen Isotopes. Figure 1 The oxygen three-isotope diagram for important end-member compositions that contributed
among the planetary compositions. The inserted figure displays the O-isotope compositions among the bulk meteorites. Note
that the terrestrial fractionation “line” (TFL) is precisely a curve when plotted over a large range of d values. Refer to the main
text for other notations

At a wavelength of 91–108 nm, CO shows narrow and
isotopically separated absorption bands leading to its pho-
todissociation through the predissociation mechanism. In
a gas medium irradiated by an ultraviolet light, the wave-
lengths which correspond to the absorption bands for
12 16
C O, the dominantly major isotopologue of CO, are
more quickly attenuated than those for other minor
isotopologues. The CO self-shielding is considered to be
responsible for the carbon and oxygen isotope fraction-
ations in molecular clouds (Visser et al. 2009). Many
workers attempted to apply the self-shielding effect to
the early solar nebula. First, Kitamura and Shimizu
(1983) proposed that 17,18O-rich O atoms derived from
the self-shielding effect in the photodissociation of O2,
exchanged with mineral grains upon shock heating, may
lead to 17,18O-enriched planetary solid materials. How-
ever, Navon and Wasserburg (1985) pointed out that the
17,18
O-enrichment produced by the self-shielding will be
rapidly erased by the reaction between the O atom and O2.
Clayton (2002) proposed that CO self-shielding may occur
at the inner edge of the solar nebula, in the proximity of the
Sun. Later models assume that the CO self-shielding
occurred at the cold environment, at the envelope of the
outer solar nebula (Lyons and Young 2005) or in the paren-
tal molecular cloud, part of which later evolved to the solar
nebula (Yurimoto and Kuramoto 2004). In these CO self-
shielding models, the 17,18O-rich isotope signature pro-
duced by the self-shielding is assumed to be transmitted to
H2O, and finally trapped by the silicate minerals.
We may test these O isotope fractionation models
by the knowledge of several important end-member
components. The most important one is the solar com-
position. The solar composition here refers to the compo-
sition of the solar gas at the surface of the Sun. The
temperature at the solar surface is low enough to prevent
occurrence of any nuclear reactions to alter the O-isotope
composition. Also, the mixing of masses between the
surface and the solar core is considered to be negligibly
small. Therefore, the composition of the gas at the
solar surface is considered to preserve the proto-solar

composition, representing the bulk composition of the
solar nebula. The MIF models assume that the solar
O-isotope composition is the same with the terrestrial
composition, whereas the CO self-shielding models
assume a 16O-rich solar composition. Hashizume and
Chaussidon (2005) discovered at the surface of several
metallic grains contained in lunar soil samples, irradiated
by the solar wind at the lunar surface since 1–2 Gyr ago, an
oxygen component enriched in 16O by >2% relative to
the terrestrial composition. They assigned the observed
16
O-rich oxygen to the solar-wind composition, conclud-
ing that the bulk solar nebula was 16O-rich. However, the
situation was complicated by the finding of a distinct
O-isotope composition at the surface of metallic grains
from a different lunar soil sample, where O was enriched
in 17,18O by 3% relative to the terrestrial composition
(Ireland et al. 2006). Hashizume and Chaussidon (2009)
confirmed existence of both the 16O- and 17,18O-rich com-
ponents among the lunar metallic grains, but concluding
that the 17,18O-rich component seems to be a nonsolar
component. This conclusion came from their observation
that the 17,18O-rich component was more than an order of
magnitude too abundant to assign it to the solar-wind
composition, whose amount was estimated from the
implanted dose of solar noble gases to the lunar soil grains.
This debate may come to the end in the near-future after
the final result for the O-isotope determination of the
solar-wind component (McKeegan et al. 2010) captured
by the GENESIS space mission. The space-probe circled
the Sun for several years to accumulate solar wind
on several substrates, which were returned to Earth
in 2004.
The other important end-member compositions,
which may serve as tests for several models, are the pri-
mordial compositions of water ice and organic matter.
The models by Yurimoto and Kuramoto (2004) or Lyons
and Young (2005) predict that water ice is the carrier of the
17,18
O-rich isotope signature, connecting the 17,18O-rich
O atoms, first produced by the CO self-shielding in the gas
medium, and the rocky materials at the mid-plane of the
solar-nebula, which finally trapped the isotope signature.
Sakamoto et al. (2007) discovered in a primitive carbona-
ceous chondrite Acfer094, a poorly characterized iron-
oxide mineral with O-isotope compositions as high as
d17O d18O 180‰. They explained that the iron-
oxide mineral was produced by reactions between primor-
dial water and the metallic Fe-Ni or iron-sulfides, and
argued that their observation demonstrates the role of
water ice as a carrier of the O-isotope signature produced
by the CO self-shielding. Hashizume et al. (2011) recently
discovered that an Antarctic carbonaceous chondrite
Oxygen Isotopes O 1199
contains pieces of organic particles that display striking
enrichments in the 17O and 18O at almost paralleled
degrees, by as high as 530‰ relative to the terrestrial
composition. The oxygen isotope anomalies were associ-
ated with carbon isotope anomalies, roughly with
a relationship of d17O 1.05  d18O 0.5  d13C.
By the CO self-shielding, the enrichment of 13C is
indeed expected to be associated with the 17,18O enrich-
ment. However, the 13C-rich signature produced among
the photodissociated products is considered to be
compromised by a competing reaction. The ion molecule
reaction (12CO + 13C+ ↔ 12C+ + 13CO + 35 K) is partic-
ularly active at low temperatures, whose isotope effect is
considered to exceed the effect by the self-shielding below
a gas temperature of 60 K (Visser et al. 2009). The positive
correlation between the d17,18O and d13C observed among
the organics suggests that the temperature where the
CO photodissociation occurred was higher than the
one for the typical molecular cloud core (10 K), which
is the place proposed by Yurimoto and Kuramoto (2004)
where the CO self-shielding might have occurred.
Hashizume et al. (2011) argued that the organic formation
associated with the 17,18O-enrichment probably occurred
at the envelope of the solar nebula, a slightly warm
(>40 K) and dense gas medium intensively illuminated
by the proto-Sun.
Future Directions
The non-mass-dependent O-isotope anomaly, broadly O
observed among the solid planetary materials, was origi-
nally considered to represent the origins or formation
processes of a subset of the rocky planetary materials.
However, recent studies suggest that the O-isotope anom-
aly is deeply connected to the origins of the low-
temperature condensates, that is, the volatiles, represented
by the water ice and the organic matter, which are critically
important materials in astrobiology. It is important to
summarize the two advantages of O-isotope over other
light element (H, C and N) stable isotopes as a tracer.
(1) Oxygen has three isotopes. We may accurately discrim-
inate the original mass-independent isotope signature
from the mass-dependent fractionations that may occur
by miscellaneous effects. (2) Oxygen is the major constit-
uent in both low- and high-temperature condensates. The
oxygen isotope may become one of the most important
tools to trace the entire history of the important ingredi-
ents for a habitable planet: from their origins; their distri-
bution processes in the solar nebula; accretion to planets/
asteroids; and the processes within the planets, namely, the
rock–water–organics interactions, possibly including the
life-formation process.
1200 O Oxygen Respiration

See also Visser R, van Dishoeck EF, Black JH (2009) The photodissociation and
chemistry of CO isotopologues: applications to interstellar clouds
▶ Fractionation, Mass Independent and Dependent
and circumstellar disks. Astron Astrophys 503:323–343
▶ Interstellar Chemical Processes Young ED, Russell SS (1998) Oxygen reservoirs in the early solar nebula
▶ Isotopic Fractionation (Interstellar Medium) inferred from an Allende CAI. Science 282:452–455
▶ Isotopic Fractionation (Planetary Process) Yurimoto H, Kuramoto K (2004) Molecular cloud origin for the oxygen
▶ Isotopolog isotope heterogeneity in the solar system. Science 305:1763–1766
▶ Photochemistry
▶ Photodissociation Regions
▶ Protoplanetary Disk, Chemistry
▶ Stellar Nucleosynthesis Oxygen Respiration
▶ Aerobic Respiration
References and Further Reading
Clayton RN (1993) Oxygen isotopes in meteorites. Annu Rev Earth Planet
Sci 21:115–149
Clayton RN (2002) Self-shielding in the solar nebula. Nature 415:860–861 Oxygenase
Clayton RN (2005) Oxygen isotopes in meteorites. In: Davis AM (ed)
Meteorites, comets and planets, vol 1, Treatise on geochemistry. Definition
Elesevier-Pergamon, Oxford, pp 129–142 Oxygenases catalyze the transference of oxygen from
Clayton RN, Grossman L, Mayeda T (1973) A component of primitive
nuclear composition in carbonaceous meteorites. Science 182:
molecular oxygen to an oxidized substrate. They are
485–488 a group of ▶ enzymes that form a class of oxydoreductases
Hashizume K, Chaussidon M (2005) A non-terrestrial 16O-rich isotopic (enzymes that catalyze the transference of electrons from
composition for the protosolar nebula. Nature 434:619–622 one molecule (called reductant) to another (the oxidant)).
Hashizume K, Chaussidon M (2009) Two oxygen isotopic components There are two classes of oxygenases: monooxygenases,
with extra-selenial origins observed among lunar metallic grains – in
search for the solar wind component. Geochim Cosmochim Acta
which transfer an oxygen atom from molecular oxygen
73:3038–3054 to the substrate and reduce the other atom to water, and
Hashizume K, Takahata N, Naraoka H, Sano Y (2011) Extreme oxygen dioxygenases, which are able to transfer both oxygen
isotope anomaly with a solar origin detected in meteoritic organics. atoms to the substrate.
Nat Geosci (in press)
Ireland Isotopic enhancements of 17O and 18O from solar wind particles
in the lunar regolith. Nature 440:776–778
See also
Kitamura Y, Shimizu M (1983) Oxygen isotopic anomaly and solar ▶ Enzyme
nebular photochemistry. Moon Planet 29:199–202 ▶ Oxidation
Lyons JR, Young ED (2005) CO self-shielding as the origin of
oxygen isotope anomalies in the early solar nebula. Nature
435:317–320
Marcus RA (2004) Mass-independent isotope effect in the earliest
processed solids in the solar system: a possible mechanism. J Chem Oxygenation
Phys 121:8201–8211
McKeegan KD, Kallio APA, Heber VS, Jarzebinski G, Mao PH, Coath CD, ▶ Oxygenation of the Earth’s Atmosphere
Kunihiro T, Wiens R, Allton J, Burnett DS (2010) Genesis SiC
concentrator sample traverse: confirmation of 16O-depletion of ter-
restrial oxygen. Lunar Planet Sci 41:2589, CD-ROM
Navon O, Wasserburg GJ (1985) Self-shielding in O2 – a possible expla-
nation for oxygen isotope anomalies in meteorites? Earth Planet Sci Oxygenation of the Earth’s
Lett 73:1–16 Atmosphere
Nguyen AN, Stadermann FJ, Zinner E, Stroud RM, Alexander C, O’D M,
Nittler LR (2007) Characterization of presolar silicate and oxide
grains in primitive carbonaceous chondrites. Astrophys J 656:
DAVID C. CATLING
1223–1240 Department of Earth and Space Sciences, University of
Sakamoto N, Seto Y, Itoh S, Kuramoto K, Fujino K, Nagashima K, Krot Washington, Seattle, WA, USA
AN, Yurimoto H (2007) Remnants of the early solar system water
enriched in heavy oxygen isotopes. Science 317:231–233
Thiemens MH (1999) Mass-independent isotope effects in planetary
atmospheres and the early solar system. Science 283:341–345
Synonyms
van Dishoeck EF, Black JH (1988) The photodissociation and chemistry of Atmospheric redox change; Earth’s atmosphere, oxygena-
interstellar CO. Astrophys J 334:771–802 tion; Great oxidation event; Oxygenation; Rise of oxygen

Keywords
Great oxidation event, rise of oxygen, oxygenation
Definition
Earth’s oxygenation is an increase in the concentration of
atmospheric molecular oxygen (O2) from levels of less
than 1 ppmv before 2.45 Ga to 21% by volume today.
Larger amounts of atmospheric oxygen became possible
because of shifts in the competition between the produc-
tion of oxygen derived from ▶ photosynthesis and the rate
of consumption of oxygen by different geological pro-
cesses. Evidence from ancient rocks suggests that oxygen-
ation happened in steps, with a first rise of O2 at 2.45–2.32
Ga and a second around 0.75–0.58 Ga. The latter increase
was a precursor to the appearance of macroscopic animals.
Overview
The present atmosphere contains (by volume) 78.05% N2,
20.95% O2, 0.93% Ar, 0.038% CO2, and various trace
gases. With the exception of argon, the concentrations of
all of major gases are biologically modulated. Oxygen, in
particular, is almost solely biogenic because it has no
significant abiotic source. Consequently, before life
existed, the atmospheric partial pressure of O2 is estimated
to have been <1013 bars. The importance of O2 cannot
be overstated: without sufficient O2, the Earth would have
? Cyanobacteria
100%
10%
1%
Atmospheric oxygen
0.1%
0.01%
0.001%
1 ppm
0.1 ppt
4.4 3.2 3.0 2.8 2.6 2.4 2.2 2.0 1.8 1.1 0.8 0.6 0.4 0.2 0.0
Time before present (Ga)
Oxygenation of the Earth’s Atmosphere. Figure 1 A schematic history of atmospheric oxygen according to geologic
constraints discussed in the text
Oxygenation of the Earth’s Atmosphere O 1201
no large multicellular plants, no land animals, and no
humans or consciousness. A particular type of photosyn-
thesis, ▶ oxygenic photosynthesis, is responsible for sup-
plying O2 to the atmosphere. Jan Ingenhousz, a Dutch
physician and botanist, discovered oxygenic photosynthe-
sis in 1779, by showing that when green plants are exposed
to sunlight they produce oxygen. In the nineteenth cen-
tury, many single-celled photosynthesizers were identified
in addition to plants, which included algae and organisms
that were initially called blue-green algae. Like other algae,
the blue-green ones were assumed to have the same cellu-
lar structure as plant cells, as a matter of definition. How-
ever, in the 1970s, blue-green algae were recognized as
bacteria and were reclassified as cyanobacteria through
the efforts of Roger Stanier, a Canadian microbiologist.
Our modern description of oxygenic photosynthesis is
a process by which green plants, algae, and cyanobacteria
split molecules of water into hydrogen and oxygen. The
hydrogen is used inside cells to chemically reduce carbon
dioxide to organic matter, while the oxygen is released as
O2 waste gas.
In the twentieth century, geochemistry uncovered the
history of Earth’s atmospheric O2 levels, showing that
for roughly the first half of Earth history, up until about
2.4 Ga, O2 concentrations were less than 1 ppmv (parts per
million by volume) (Fig. 1). The first person to describe
O
Visible fossils of algae
Animals
1202 O Oxygenation of the Earth’s Atmosphere
evidence that O2 levels were significantly different on the
early Earth was the geologist Alexander MacGregor. In the
1920s, MacGregor examined Archean (pre-2.5 Ga) sedi-
mentary rocks in modern-day Zimbabwe and deduced
that they formed in an atmosphere devoid of O2 because
they lacked the reddish-colored, oxidized iron minerals
that are ubiquitous in analogous modern sediments.
MacGregor’s conclusion greatly influenced Preston
Cloud, an American geologist, who further investigated
the geologic record for information on past atmospheric
O2. Cloud also championed a suggestion originally made
in 1959 by the marine biologist John Nursall that the
relatively late appearance of macroscopic fossils animals
in the geologic record reflected the slowness in reaching
a sufficiently O2-rich atmosphere.
Modern geochemical research shows that an initial
transition to an oxygenated atmosphere occurred in the
period 2.45–2.32 Ga, but the O2 levels attained were prob-
ably only 0.2–2% by volume and apparently insufficient
for supporting macroscopic animals. This first “rise of
oxygen” is also called the Great Oxidation Event (GOE).
Although much less than today’s 21% by volume, the
post-GOE O2 levels were enough to dominate the redox
chemistry of the atmosphere. Such O2 levels were also
ample for producing a stratospheric ozone (O3) layer,
which ever since has protected life on the Earth’s surface
from biologically harmful ultraviolet radiation. The first
fossils visible to the naked eye occur after the GOE at
1.87 Ga and are thought to represent a seaweed, Grypania
spiralis. In a second increase in O2, over the period 750–
580 Ma, O2 rose to levels of at least tens of percents of
modern, that is, a concentration between 2% and present
levels. The first animal fossils appear in the Ediacaran Period
(635–542 Ma), with the oldest soft-bodied fossils of Edia-
caran fauna at 575–565 Ma. While there is a consensus that
sufficient O2 was a necessary physiological precursor to
animal life, there remains considerable debate among
paleontologists about whether an increase in O2 caused
large animals to appear or whether other evolutionary and
ecological factors were crucial.
Explaining why O2 increased is more elusive than it
might first appear. We might suppose that when
cyanobacteria evolved, O2 would have accumulated in
the atmosphere. But the consensus interpretation of geo-
chemical evidence (discussed below) is that cyanobacteria
produced O2 for several hundred million years or
more before 2.4 Ga, while atmospheric O2 levels remained
<1 ppmv. A pertinent consideration is that the Earth’s
overall chemical composition is reducing, so that reactions
of O2 with geothermal gases in the atmosphere or certain
cations in the ocean (e.g., Fe2+) in an initially reducing
environment must have acted as a powerful chemical
buffer opposing the accumulation of atmospheric O2.
Consequently, we need to carefully consider the losses of
O2 as well as its source for understanding past changes in
O2 levels. In this regard, the modern oxygen cycle helps to
elucidate some of the basic concepts of source and sink
fluxes before discussing how sources and sinks were prob-
ably different on the ancient Earth.
The O2 Balance Sheet: Gains and Losses
The theoretical subtleties of how photosynthesis can influ-
ence long-term atmospheric O2 levels were not fully
appreciated until the 1970s, in spite of the fact that
a French geologist, Jacques Ebelmen, correctly described
the key ideas in 1845. Ebelmen implicitly realized that
photosynthesis is nearly a zero-sum activity. Oxygenic
photosynthesis can be summarized by the following sche-
matic equation:
CO2 þH2 O $ CH2 OþO2 ð1Þ
where “CH2O” represents the average stoichiometry of
organic matter. Oxygen consumption by respiration or
decay of organic matter reverses Eq. 1, and produces no
net oxygen on timescales greater than about 102 years.
Oxygen only escapes respiration of oxidative decay on
geologic timescales because a tiny fraction (0.1–0.2%) of
organic carbon is buried, mainly in marine sediments,
which segregates the oxygen from the organic matter.
Some organic matter is also used in biogeochemical pro-
cesses to make pyrite (FeS2) from seawater sulfate ions
(SO42(aq)) as follows:
Oxygenic photosynthesis : CO2 þH2 O ¼ CH2 O þ O2 15
Anaerobic decomposition : 15CH2 O þ 8SO4 2 þ 2Fe2 O3
þ 16Hþ ¼ 4FeS2 þ 15CO2 þ 23H2 O

Net reaction : 8SO4 2 þ 2Fe2 O3 þ 16Hþ
¼ 15O2 þ 4FeS2 ðburiedÞ þ 8H2 O
ð2Þ
Consequently, pyrite burial also liberates oxygen into
the atmosphere or ocean. Remarkably, Ebelmen correctly
identified both organic carbon burial and pyrite burial as
the long-term sources of atmospheric oxygen. Unfortu-
nately, his ideas went unnoticed and the same concepts
had to be reinvented.
While it is useful to think of the burial of organic
carbon and pyrite as a “net source” of O2, it is important
to appreciate that this “net” O2 is lost to numerous other
sinks for O2 besides the reversal of eq. [1]. The geological
sinks and sources of O2 are shown in Fig. 2. By reviewing
 Oxygenation of the Earth’s Atmosphere O 1203

Escape of hydrogen to space

Subaerial volcanic gases

Continental
weathering Oxygen, O2

Metamorphic
gases

Submarine Submarine
Burial of organic volcanic weathering
carbon or pyrite gases

Oxygenation of the Earth’s Atmosphere. Figure 2 Schematic diagram of source and sink fluxes of oxygen on geologic
timescales

Oxygenation of the Earth’s Atmosphere. Table 1 Modern source fluxes of O2 (Holland 2002)

Oxygen fluxes 1012 mol O2 year-1 Effect on O2

Organic carbon burial flux 10.0  3.3 Production
Pyrite (FeS2) burial flux 7.8  4.0 Production
Sulfate burial flux (0.3  0.1) (Loss)
Reduced iron burial flux 0.9  0.4 Production O
Total rate of oxygen production  (18.4  7.8)  10 12 -1
mol O2 year

the average composition of new sediments, American
geochemist Heinrich Holland has estimated that burial
fluxes of organic carbon and pyrite produce oxygen at
rates of 10.0  3.3 Tmol O2 year1 and 7.8  4.0 Tmol
O2 year1, respectively, where 1 Tmol = 1012 mol (Holland
2002). The reduction of oxidized iron from its +3 oxida-
tion state to the +2 state (2Fe2O3 = 4FeO + O2) and
subsequent burial also adds a minor flux of O2, whereas
the burial of sulfate minerals in sediments removes O2.
Summing these burial fluxes, the total O2 net source is
18.4  7.8 Tmol O2 year1 (Table 1). Dividing the current
amount of oxygen in the atmosphere-ocean reservoir
(3.78  1019 mol O2) by the source flux (1.8  1013 mol
O2 year1) gives 2.1 million years for the average
amount of time an O2 molecule spends in today’s atmo-
sphere-ocean system. Consequently, over such timescales,
the net source of O2 must be closely balanced by losses or
O2 levels would wildly fluctuate. We know that
centimeter- to meter-sized animals, which require O2 to
grow and breathe, have been continuously present in the
Phanerozoic eon (542 Ma to present) so that concentra-
tions of atmospheric O2 must have been within 0.2  0.1
bar. The O2 source is balanced by O2 losses that include
reactions in the ocean with dissolved minerals and gases
from hot seafloor vents, and reactions of O2 dissolved in
rainwater with continental minerals, principally those
containing elemental carbon, sulfide, and ferrous iron
(Fig. 1). Gases that consume O2 in the atmosphere or
ocean are CH4, H2, CO, SO2, and H2S. These are released
either from hot rocks that melt (volcanism) or hot rocks
that do not melt (metamorphism). Estimates of their
fluxes are shown in Table 2. The O2 sink from submarine
and subaerial volcanic gases is 2.3 Tmol O2 year1,
metamorphic gases consumes 3.1 Tmol O2 year1, and
subduction of oxidized seafloor causes a loss of 0.2 Tmol
O2 year1. The sink due to continental weathering is
1204 O Oxygenation of the Earth’s Atmosphere

Oxygenation of the Earth’s Atmosphere. Table 2 Modern sink fluxes of O2 (Holland 2002; Catling and Kasting 2012)

Type of flux 1012 mol O2 year1 Effect

Continental oxidative weathering (15.5  6.7) (Loss)
Surface volcanic gases
H2 (0.5  0.3) (Loss)
CO (0.05  0.03) (Loss)
H2S (0.06  0.03) (Loss)
SO2 (0.9  0.3) (Loss)
1
Total sink from surface volcanic gases   (1.5  0.7)  10
12
mol O2 year
Submarine volcanic gases
H2 (0.05  0.03) (Loss)
H2S (0.7  0.4) (Loss)
CH4 (axial) (0.02  0.01) (Loss)
CH4 (off-axis) (0.5  0.3) (Loss)
1
Total sink from submarine volcanic gases   (0.8  0.7)  10 12
mol O2 year
Surface metamorphic gases
CH4 (abiotic) 0.6 (Loss)
CH4 (thermal breakdown of organics) 2.5 (Loss)
Total sink from metamorphic gases 3.1  1012 mol O2 year1
Seafloor oxidation and subduction
Fe2+ conversion to magnetite 0.2 (Loss)
Total rate of oxygen loss  (21.1  8.1)  1012 mol O2 year1

15.5  6.7 O2 year1, which is estimated from the com-
position of average rock undergoing weathering (Table 2).
Thus, within the large 40% uncertainty of the data, the
source flux of O2 from organic carbon and pyrite burial of
18.4  7.8 Tmol O2 year1 is matched by the total sink flux
of 21.1  8.1 Tmol O2 year1. In the past, changes in the
relative proportions of source and sink fluxes evidently
caused O2 levels to change.
Evidence for Earth’s Oxygenation
The Great Oxidation Event at 2.45–2.32 Ga occurred in
the Paleoproterozoic era (2.5–1.6 Ga), the first of three
eras within the Proterozoic eon (2.5–0.542 Ga), which
follows the Archean eon (before 2.5 Ga). After the
Paleoproterozoic, the Mesoproterozoic (1.6–1.0 Ga) and
the Neoproterozoic (1.0 Ga–542 Ma) eras follow. The
second rise of O2 (750–580 Ma) occurred in the
Neoproterozoic. We can tell that O2 levels changed by
studying at the chemistry of rocks from these eras. The
Great Oxidation Event (GOE) produced binary geochem-
ical changes in the Paleoproterozoic because the atmo-
sphere changed the oxidation states (from weakly
reducing to oxidizing), whereas the Neoproterozoic
changes are subtler because an oxic atmosphere merely
became more oxygenated. Let us consider evidence for the
GOE and Neoproterozoic increase of O2 in turn.
Evidence for the Great Oxidation Event
(GOE)
The different oxidation state of continental minerals that
were formed before and after the GOE records a change
from an atmosphere essentially devoid of O2 to one redox
dominated by O2. The presence or absence of O2 changes
the chemistry of minerals that are formed because O2
dissolves in rainwater and reacts with minerals. Three
continental indicators are red beds, paleosols, and detrital
grains (Holland 1984). Red beds are riverbanks, deserts,
and floodplains with a reddish pigmentation that arises
when an iron oxide coating on mineral grains is produced
from the rusting of iron minerals. Before the GOE, no red
beds are found in the geologic record, indicating a lack of
O2, whereas afterward red beds are common. Paleosols are
lithified soils. Before the GOE, paleosols show a loss of
iron, which is consistent with leaching by ancient anoxic
rainwater, whereas after 2.2 Ga, iron is not leached. Iron
will be flushed through a soil in the form of soluble ferrous
iron ions (Fe2+) if rainwater has little dissolved O2 but if
the air contains >0.2% O2, iron will be oxidized into

insoluble and immobile oxides such as ▶ hematite
(Fe2O3). Cerium is another redox-sensitive element that
has also been examined in paleosols. Cerium oxidizes
from Ce3+ to Ce4+ to form cerianite (CeO2), so the pres-
ence of Ce3+-rich minerals in paleosols from the Archean
implies an early anoxic atmosphere. Detrital grains in
sediments are those that are never completely dissolved
during the process of weathering. Rounded detrital grains
from sediments deposited by turbulent Archean rivers
commonly contain minerals that would only survive at
very low levels of O2. Detrital grains of pyrite (FeS2),
uraninite (UO2), and siderite (FeCO3) place bounds on
the concentration of Archean O2 of roughly <2%, <0.2%,
and <0.02%, respectively, compared to today’s 21% O2.
In modern oxygenated waters, uraninite dissolves to
form soluble U6+ ions, pyrite oxidizes to soluble sulfate
(SO42) and hematite, and siderite rusts to produce insol-
uble hematite.
The chemistry of seawater also changed after the GOE,
with increased levels of sulfate and much decreased levels
of redox-sensitive elements such as manganese and iron
that are insoluble when oxidized. Marine sulfur isotopes
are the data that indicate an increase in sulfate concentra-
tions. Today, sulfate-reducing bacteria produce most of
the sulfide in marine sediments. These bacteria reduce
32
SO42 in preference to 34SO42 so that bacterial sulfide
is enriched in 32S. However, this fractionation ceases in
waters with <0.2 mM sulfate concentration. Archean sul-
fides have 34S/32S ratios close to an unfractionated mantle
value, implying Archean oceans with <0.2 mM sulfate,
compared to 28.9 mM in today’s surface seawater. Rivers
ultimately supply sulfate to the ocean by washing sulfate
off the land where it is produced by the oxidation of
sulfides by oxygenated rainwater. Consequently, the low
concentration of seawater sulfate before the GOE is con-
sistent with low atmospheric O2. By 2.3–2.2 Ga, sulfides
with significant 34S depletions formed ubiquitously in the
ocean, reflecting increased sulfate and the rise of O2. When
O2 rose, it also oxidized and titrated manganese out of the
ocean, producing the world’s largest manganese deposit
found in the 2.2 Ga Hotazel Formation in South Africa.
The presence of ▶ banded iron formations (BIFs) pro-
vides evidence that the deep ocean was devoid of O2 in
the Archean, unlike today. By definition, BIFs are lami-
Oxygenation of the Earth’s Atmosphere O 1205
was (microbially) oxidized and precipitated as iron min-
erals in the BIFs. In the Paleoproterozoic, BIFs decline in
abundance and disappear after 1.8 Ga. Chromium (Cr)
isotopes in the BIFs also record changes in atmospheric
O2. When insoluble Cr3+ is oxidized to soluble Cr6+ dur-
ing oxidative weathering, the heavy isotope 53Cr is prefer-
entially oxidized relative to 52Cr. After transport from
rivers to the ocean, the Cr isotopic fractionation is pre-
served in ocean sediments. An increase in the abundance
of 53Cr after 2.4 Ga compared to Archean samples further
confirms the GOE.
The strongest evidence for the GOE is a large amount
of mass-independent fractionation (MIF) of ▶ sulfur iso-
topes in sedimentary rocks older than 2.45 Ga compared
to little or no sulfur isotope MIF in rocks younger than
2.32 Ga. In modern rocks, 32S, 33S, and 34S obey “mass-
dependent” fractionation, in which the difference in abun-
dance between 33S and 32S is approximately half that
between 34S and 32S, that is, a linear relationship with
mass difference. Most processes, such as microbial sulfate
reduction, produce mass-dependent isotope fraction-
ation. In contrast, mass-independent fractionation in sul-
fur isotopes is where the relative abundance of different
isotopes deviates from linear, mass-dependent fraction-
ation. Laboratory studies show that large MIF is produced
exclusively from photochemical reactions, such as the
photolysis of SO2. Three conditions are needed for the
production and preservation of the sulfur isotope MIF: O2
levels below about 1 ppmv, sufficient sulfur gas in the O
atmosphere, and relatively large concentrations of reduc-
ing gases. In a high-O2 atm, sulfur gases are rapidly oxi-
dized and rained out as dissolved sulfate. But in the
absence of O2 and in the presence of reducing gases such
as 102–103 ppmv methane, sulfur exits the atmosphere as
sulfide, elemental sulfur, or sulfate, which allows the mass-
independent fractionation produced by anoxic photo-
chemistry to be segregated and preserved in different
sedimentary minerals. In summary, the sulfur isotope
MIF provides evidence for an Archean atmosphere with
both low O2 and high methane.
Donald Canfield (of the University of Southern
Denmark) has proposed that a significant fraction of the
deep ocean remained sulfidic rather than oxygenated dur-
ing much of the Proterozoic, after 1.8 Ga (Canfield 2005;
nated marine sedimentary rocks containing  15 wt%
iron, with alternating iron-rich and iron-poor layers. The
iron mainly originated from hydrothermal sources in the
deep ocean, such as midocean ridges. Today, iron is oxi-
dized in the deep ocean and deposited on the flanks of the
ridges. But in the anoxic depths of Archean oceans, ferrous
iron was transported to the continental shelves where it
Lyons et al. 2009). In this model, BIFs declined because
increased sulfate in the surface ocean caused sulfate-
reducing bacteria to provide a flux of sulfide to the deep
ocean, which scavenged ferrous iron into insoluble iron
sulfides. This is an alternative to a model where the disap-
pearance of BIFs at 1.8 Ga is explained by the oxygenation
of the deep ocean and the formation of insoluble iron
1206 O Oxygenation of the Earth’s Atmosphere
oxides (Holland 2006). In today’s ocean, <0.5% of the
seafloor is euxinic, meaning oxygen free with H2S. Molyb-
denum (Mo) isotopes are useful indicators of oxygenation
and euxinia because oxic uptake of Mo on marine solids
preferentially removes 95Mo relative to 98Mo, leaving the
ocean isotopically heavy. In contrast, in times of low
oxygenation, marine Mo is isotopically light. In highly
euxinic deep waters, Mo is scavenged efficiently and
tends to capture the Mo isotope composition of seawater.
Rocks that are 95Mo-enriched from the 1.7–1.4 Ga
MacArthur Basin in Australia suggest that the deep ocean
in the mid-Proterozoic seafloor was commonly oxygen-
poor, with a greater proportion of euxinia than today. The
data suggest that the euxinia was probably only several
percent of the ocean’s area, but this is enough to have had
a significant effect in depleting trace metals in the
Mesoproterozoic Ocean, which may have impacted marine
ecology for a billion years until the Neoproterozoic.
Evidence for a Neoproterozoic “Second Rise”
in Oxygen
Sulfur and chromium isotopes in marine sediments indi-
cate that O2 increased in the late Neoproterozoic. At
this time, marine sedimentary sulfides occur with
32
S-enrichment exceeding the isotope discrimination
of sulfate-reducing bacteria. The interpretation is that
isotopically light sulfide was re-oxidized at the sediment-
water interface and cyclically re-reduced by sulfur-
disproportionating bacteria, which increased the isotope
fractionation. The greater sulfur isotope fractionation is
therefore related to oxygenation of the near-shore seafloor.
Canfield (2005) has estimated that the oxygenation
required atmospheric O2 concentrations in excess of
1–4%. A large increase in the abundance of marine 53Cr
relative to 52Cr after 750 Ma also confirms an increase in
atmospheric O2. Oxygen would have mobilized greater
amounts of chromium during continental oxidative
weathering, which is the mechanism of isotopic fraction-
ation mentioned earlier. Fossil evidence for animals
greater than a few millimeters or more in size also suggests
that O2 concentrations exceeded 2%, which is the mini-
mum level that can support aerobic metabolism in such
animals via O2 supplied by diffusion.
When Did Oxygenic Photosynthesis Evolve?
In order to understand the early history of atmospheric
O2, we need to examine the evidence for the advent of
oxygenic photosynthesis. In particular, the geologic record
suggests that oxygenic photosynthesis arose 300 Ma or
more before the GOE. This timing is inferred from
a variety of proxies including biomarkers, nonmarine
stromatolites, redox-sensitive metals, and sulfur isotopes,
which we discuss in turn.
Biomarkers are recognizable derivatives of biological
molecules, which are usually in the form of certain hydro-
carbons found in kerogen (noncrystalline sedimentary
organic matter) or oil. Chemical changes induced when
sediments convert into rock strip biological molecules of
functional groups but can often leave a distinctive molec-
ular backbone, in a way that is analogous to the more
familiar macroscopic process that removes flesh to leave
fossilized skeletons that we can associate with particular
life forms. Biomarkers from the Archean have been the
subject of controversy because of the possibility of con-
tamination from modern molecules or fluid migration
over geologic time and uncertainties regarding their
specificity for particular organisms and metabolisms.
Recent advances in ultraclean drilling and the ability to
analyze oil-bearing fluid inclusions help to address con-
cerns about contamination. Biomarkers for cyanobacteria
are 2-alpha-methylhopanes, which are derived from com-
pounds in cell membranes. Long chain varieties of these
particular molecules (i.e., >C31) occur very rarely outside
the cyanobacteria. In addition, steranes derive from ste-
rols, which are found in the membranes of eukaryotes and
synthesized in a pathway that requires free molecular
oxygen. For example, 11 molecules of O2 are required to
produce one molecule of cholesterol. No bacteria are
known to synthesize C26–C30 sterols and archaea do not
synthesize sterols at all. The simultaneous co-detection of
long chain (>C31) 2-alpha-methylhopanes with eukary-
otic sterols is thus a reasonable biosignature for oxygenic
photosynthesis. Such a combination of biomarkers has
been found in fluid inclusions in the 2.45 Ga Matinenda
Formation at Elliot Lake, Canada, which consists of sand-
stones deposited prior to the GOE (Dutkiewicz et al.
2006). Also, meticulous measurements of biomarkers
from 2.72 to 2.56 Ga rocks in the Hamersley Province
of northwest Australia (Eigenbrode et al. 2008) and
2.67–2.46 Ga rocks from the Transvaal of South Africa
(Waldbauer et al. 2009) suggest the presence of microbial
ecosystems that produced O2 as well as consumed it.
Stromatolites (laminated rocks formed by microbes)
from a 2.72 Ga lake in the Tumbiana Formation in north-
west Australia possibly confirm the presence of O2
production. Geologic evidence shows that the lake
lacked sulfate and hydrothermal reductants needed
for non-oxygen-producing photosynthesis. Consequently,
a process of elimination suggests that microbial
communities using oxygenic photosynthesis may
have constructed these particular stromatolites (Sakurai
et al. 2005).

Furthermore, the mobility of redox-sensitive metals
indicates the presence of some O2 before the GOE, and
therefore the presence of oxygenic photosynthesis. Molyb-
denum (Mo), rhenium (Re), and chromium (Cr) are
soluble only when oxidized. Consequently, a spike in Mo
and Re concentrations found in 2.5 Ga sedimentary rocks
in the Hamersley district of northwest Australia suggests
that a transient “whiff ” of O2 occurred in the atmosphere
about 100 million years prior to the GOE (Anbar et al.
2007). The whiff is confirmed by contemporaneous evi-
dence of sulfur isotopic fractionation indicating a spike of
sulfate. Enrichments of 53Cr relative to 52Cr (which are
induced by oxidative weathering) are also found in banded
iron formations at low levels for at least 300 million years
prior to the GOE, which is consistent with the biological
production of O2 (Frei et al. 2009). In summary, multiple
lines of evidence suggest that oxygenic photosynthesis
existed at least 0.3 billion years before the GOE.
Theories for Oxygenation
At present, a framework has been developed for under-
standing why the GOE occurred, but the cause of the
second rise of O2 in the Neoproterozoic remains very
obscure (Catling and Claire 2005).
The GOE happened because either O2 production
increased or O2 consumption declined. The ancient, pre-
2.45 Ga oceans were largely devoid of sulfate, so that
organic carbon burial dominated O2 production prior to
the GOE, while pyrite burial was unimportant. Conse-
quently, it has been argued that O2 rose because the rate
of organic carbon burial (O2 production) increased, either
as a pulse or long-term trend. One suggestion is that the
growth of early continental shelves promoted organic
carbon burial and oxygen production. But there are
grounds for skepticism about this proposal. Organic mat-
ter extracts the light carbon isotope, 12C, from seawater,
but marine carbon, recorded in ancient limestone,
does not significantly decrease in 12C content between
mid-Archean and Mesoproterozoic rocks. Notably, at
2.3–2.1 Ga, many marine carbonates are unusually
depleted in 12C relative to 13C, which could be explained
if there was a large pulse of burial of organic carbon, but
then the 12C content returns to its previous value. A burial
pulse would merely generate a parallel pulse of O2, given
the short residence time of O2 and not a persistently
increased level of O2. Instead, the pulse was more plausibly
an effect of increased O2 because the Paleoproterozoic
excursions in the 12C of limestones follow the rise of O2
before 2.32 Ga.
A widely accepted explanation for why little atmo-
spheric O2 accumulated in the Archean is that a large
Oxygenation of the Earth’s Atmosphere O 1207
flux of volatile and aqueous reductants, that is, chemicals
that consume oxygen, scavenged the O2. In time, the
geological sources of reductants declined to the point
where the O2 flux produced from organic carbon burial
exceeded the kinetically rapid consumption flux. At this
tipping point, oxygen flooded the atmosphere until O2
levels reached a new plateau where continental weathering
became the predominant balance for O2 production.
Various suggestions have been made for why the propor-
tion of reductants in volcanic or metamorphic outgassing
might diminish and cause such a tipping point. One
suggestion is that a change in the style of outgassing
from submarine to subaerial volcanism favored a decline
in the proportion of reductants. Another consideration is
that an anoxic atmosphere would always allow a relatively
high concentration of hydrogen-bearing reducing gases.
At high altitude, the decomposition of such gases causes
hydrogen to escape to space at a rate that is significant on
geologic timescales, which oxidizes the whole Earth and
diminishes further release of reductants. In particular, in
an anoxic atmosphere, methane reaches concentrations
hundreds of times greater than today’s 1.8 ppmv. Most
methane is ultimately produced microbially from fermen-
tation of photosynthesized organic matter. Ultraviolet
light decomposes methane in the upper atmosphere, caus-
ing it to lose hydrogen. When hydrogen escapes from the
planet, oxygen is gained irreversibly through a schematic
reaction as follows:
O
CO2 þ H2 O þ biology ! CH4 þ O2
þ photochemistry ! 4Hðlost to spaceÞ þ CO2 þ O2
ð3Þ
Evidence for this hypothesis is found in the redox
components of the crust. If photosynthesis were the only
process responsible for segregating redox components, the
number of moles of equivalent O2 in the Earth’s crust
should be balanced by equimolar reduced carbon
(Eq. 1). But instead there is excess oxygen in the crust by
a factor of 1.5–2.2 (Catling and Claire 2005; Sleep 2005).
The excess can be quantitatively reconciled with time-
integrated hydrogen escape and net oxidation expected
from a methane-rich Archean atmosphere.
Various geological and biological causes have been
suggested for the Neoproterozoic second rise of oxygen,
but the problem remains unsolved. Geological proposals
include the enhanced production of clays that adsorbed
and buried organic matter, or the assembly of
a supercontinent whose weathering added nutrients to
the sea and stimulated organic carbon burial. Also, it has
been suggested that moderately high levels of methane
1208 O Oxygenation of the Earth’s Atmosphere
throughout the Proterozoic could have promoted hydro-
gen escape and oxidized the Earth. Biological proposals
include continental weathering accelerated by lichen, and
the evolution of zooplankton with fecal pellets that
enhanced organic burial. However, presently, all of these
ideas for the cause of the second rise of O2 are, at best, just
qualitative proposals.
Conclusion and Future Directions
The atmosphere evolved from an anoxic to oxygenated state.
There are multiple lines of geochemical evidence for a rise of
O2 or Great Oxidation Event (GOE) at 2.45–2.32 Ga.
However, oxygenic photosynthesis appears to have existed
at least 0.3 billion years beforehand. A self-consistent
explanation for the delay between the origin of oxygenic
photosynthesis and oxygenation of the atmosphere is that
excess volatile reductants efficiently scavenged O2. This
sink on O2 diminished until the GOE occurred. Following
the GOE, sulfate levels increased in the ocean as a result of
a greater role for oxidative weathering in balancing O2
production from the burial of organic carbon and pyrite.
In the late Precambrian, around 750–580 Ma, a second rise
of O2 occurred, along with a further increase in marine
sulfate. This second rise appears to have been a precursor
to the appearance of macroscopic animals.
Considerable further research will no doubt further
constrain the levels of O2 through geologic time and our
understanding of the past biogeochemical cycles of
various elements – such as carbon, sulfur, and nitrogen.
Measurements of new isotopic systems, such as those of
trace metals, will no doubt bring advances. Biomarkers
also have the potential to reveal what microorganisms and
microbial processes were present in the Precambrian
biosphere. So it is likely that further study of biomarkers
will help us better understand the role of microbial
ecology in shaping the oxidation state of Earth’s ancient
atmosphere.
See also
▶ Archean Environmental Conditions
▶ Archean Traces of Life
▶ Banded Iron Formation
▶ Biomarkers
▶ Carbon Isotopes as a Geochemical Tracer
▶ Earth, Formation and Early Evolution
▶ Earth’s Atmosphere, Origin and Evolution of
▶ Great Oxygenation Event
▶ Hematite
▶ Iron Oxyhydroxides
▶ Isotope Biosignatures
▶ Nitrogen Isotopes
▶ Oceans, Origin of
▶ Oxidizing Atmosphere
▶ Oxygenic Photosynthesis
▶ Photosynthesis
▶ Pilbara Craton
▶ Redox Zonation
▶ Siderite
▶ Sulfidic Oceans
▶ Sulfur Cycle
▶ Sulfur Isotopes
▶ Transition Metals and Their Isotopes
▶ Transvaal Supergroup, South Africa
References and Further Reading
Anbar AD et al (2007) A whiff of oxygen before the great oxidation event?
Science 317:1903–1906
Canfield DE (2005) The early history of atmospheric oxygen: homage to
Robert A. Garrels. Annu Rev Earth Planet Sci 33:1–36
Catling DC, Claire MW (2005) How Earth’s atmosphere evolved to an
oxic state: a status report. Earth Planet Sci Lett 237:1–20
Catling DC, Kasting JF (2012) Atmospheric Evolution on Inhabited and
Lifeless Words. Cambridge University Press, Cambridge
Dutkiewicz A, Volk H, George SC, Ridley J, Buick R (2006) Biomarkers
from Huronian oil-bearing fluid inclusions: an uncontaminated
record of life before the Great Oxidation Event. Geology 34:437–440
Eigenbrode JL, Freeman KH, Summons RE (2008) Methylhopane
biomarker hydrocarbons in Hamersley Province sediments provide
evidence for Neoarchean aerobiosis. Earth Planet Sci Lett
273:323–331
Frei R, Gaucher C, Poulton SW, Canfield DE (2009) Fluctuations in
Precambrian atmospheric oxygenation recorded by chromium
isotopes. Nature 461:250–253
Holland HD (1984) The Chemical Evolution of the Atmosphere and
Oceans. Princeton University Press, Princeton
Holland HD (2002) Volcanic gases, black smokers, and the Great Oxida-
tion Event. Geochim Cosmochim Acta 66:3811–3826
Holland HD (2006) The oxygenation of the atmosphere and oceans.
Philos Trans R SocB-Biol Sci 361:903–915
Knoll AH (2003) Life on a Young Planet: The First Three Billion Years of
Evolution on Earth. Princeton University Press, Princeton
Lyons TW, Anbar AD, Severmann S, Scott C, Gill BC (2009) Tracking
euxinia in the ancient ocean: a multiproxy perspective and Protero-
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Oxygenic Photosynthesis
Definition
Oxygenic photosynthesis is a non-cyclic photosynthetic
electron chain where the initial electron donor is water
and, as a consequence, molecular oxygen is liberated as
a byproduct. The use of water as an ▶ electron donor
requires a photosynthetic apparatus with two reaction
centers. It is present in ▶ cyanobacteria and its evolution-
ary related eukaryotic organelles, the ▶ chloroplast.
Oxygenic ▶ photosynthesis was responsible for the trans-
formation of the Earth’s primitive anoxygenic atmosphere
to one with molecular oxygen.
See also
▶ Chlorophylls
▶ Chloroplast
▶ Cyanobacteria
▶ Electron Donor
▶ Photosynthesis
Oxygenic Phototrophic Bacteria
▶ Cyanobacteria, Diversity and Evolution of
Oxyphotobacteria
▶ Cyanobacteria, Diversity and Evolution of
Ozone
Definition
Ozone is an important trace constituent of the stratosphere
where it forms the ozone layer. It consists of three atoms of
oxygen arranged as an open triangle. Ozone is produced
in the three-body reaction O þ O2 þ M = O2 þ M,
where M represents a stabilizing molecule (N2 or O2)
and O is produced by the photochemical dissociation
of O2. Some other components (NO, Cl, CFCs) catalyze
the reaction of ozone destruction O3 þ O = 2O2.
Ozone Layer O 1209
The importance of ozone for life preservation lies in its
unique capability to block solar UV in a range particularly
harmful to organic macromolecules (230–290 nm). It is
also a greenhouse gas.
See also
▶ Fractionation
▶ Fractionation, Mass Independent and Dependent
Ozone Layer
GERDA HORNECK
German Aerospace Center (DLR), Institute of Aerospace
Medicine, Cologne, Germany
Synonyms
Stratospheric ozone
Keywords
Oxygen photochemistry, ozone, UV screening
Definition
The ▶ ozone layer refers to the ozone within the strato-
sphere of the Earth, where over 90% of the Earth’s ozone
resides. This layer absorbs the Sun’s high-energy ultravio- O
let radiation of wavelengths <290 nm. The concentration
of ozone is measured in Dobson units (DU).
History
The existence of an ozone layer in the upper regions of our
atmosphere was detected in 1913 by Charles Fabry and
Henri Buisson. In 1924, G.M.B. Dobson built the first
spectrograph to measure the seasonal variation of ozone.
From 1928 to 1958, Dobson established a worldwide net-
work of ozone monitoring stations that are operating up to
date. In his honor the “Dobson unit” was termed as
measure of the columnar density of ozone above the
Earth surface.
Overview
On Earth, the stratospheric ozone layer is found at an
altitude from 20 to 40 km. It is created when ultraviolet
radiation from the Sun strikes the oxygen molecules in the
stratosphere. By these photochemical processes oxygen
molecules (O2) are dissociated to atomic oxygen (O).
The atomic oxygen quickly combines with further oxygen
molecules to form ozone (O3). Ozone can be split again
1210 O Ozone Layer
10−17
10−18
10−19
10−20
O3 cross section/cm2
10−21
10−22
10−23
10−24
10−25
10−26
10−27
10−28
10−29
10−30
240 280 320 360
λ /nm
Ozone Layer. Figure 1 Absorption cross-section of ozone as
a function of wavelength
into molecular oxygen (O2) and atomic oxygen under the
influence of short wavelength ultraviolet radiation from
the Sun (Fig. 1). This continuing process is called ozone-
oxygen cycle.
The concentration of ozone in the stratosphere is
measured in Dobson units (DU) – named after Gordon
M.B. Dobson, who built the first instrument for measur-
ing total ozone from the ground. It determines the total
amount of ozone present in a vertical column of air above
a certain location at the surface of the Earth. It gives the
mm thickness of ozone at an atmospheric pressure of
1013 hPa and a temperature of 298 K: 1 mm of ozone is
equivalent to 100 DU. Concentrations of ozone in the
stratosphere vary naturally according to temperature,
weather, latitude, and altitude. Furthermore, aerosols
and other particles ejected by natural events such as vol-
canic eruptions can have measurable impacts on ozone
levels. Near the tropics, values typically reach about
260 DU; these values increase toward the poles.
The stratospheric ozone layer is very effective at
screening out solar UV radiation at wavelengths
<290 nm. As a result, most of the biologically harmful
part of the solar UV radiation does not reach the surface of
the Earth. Surface UVB radiation levels are highly variable
because of sun angle, cloud cover, and also because of local
effects including pollutants and surface reflections.
Decreasing ozone concentrations result in higher irradi-
ances of the energetic part (UVB) of the solar spectrum
with possible biological consequences (Häder 1997).
The Antarctic ozone hole is an area of the Antarctic strato-
sphere wherein the total ozone amount is less than
220 DU. It occurs during the Antarctic spring, from
September to early December, when over 50% of the
lower stratospheric ozone is destroyed. Over the past two
decades, the ozone hole has steadily grown in size (up to
an area of 27 million km2) and length of existence (from
August through early December).
In the Archean (3.8–2.5 billion years ago), the Earth
atmosphere was essentially anoxic and therefore it proba-
bly lacked a protective ozone column. Thus, UVB (280–
315 nm) and UVC (200–280 nm) radiation could have
penetrated to the Earth’s surface with their associated
400
biological effects. Using data obtained from the UV-
inactivation of Bacillus subtilis spores in a space experi-
ment, it was shown that without UV screen by strato-
spheric ozone the biologically effective UV doses would
be about 1,000 times higher than today (Horneck et al.
1996; Cockell and Horneck 2001). Most of the damage
would have been caused by UVC radiation in the DNA,
which is the decisive target for inactivation and mutation
induction at this UV range.
Mars has no planetary ozone layer, although regional
small concentrations of ozone may occur during northern
winter periods. As a result, most of the Martian surface
is reached by the high energetic solar UV radiation of
wavelengths >200 nm.
See also
▶ Ozone
▶ Photobiology
▶ Solar UV Radiation (Biological Effects)
▶ UV Climate
▶ UV Radiation (Biological Effects)
▶ UV Radiation Dose
References and Further Reading
Cockell CS, Horneck G (2001) The history of the UV radiation climate of
the Earth – theoretical and space-based observations. Photochem
Photobiol 73:447–451
Graedel TE, Crutzen PJ (1993) Atmospheric change: an Earth system
perspective, 2nd edn. Freeman, New York
Häder DP (ed) (1997) The effects of ozone depletion on aquatic ecosys-
tems. Academic Press, RG Landes
Horneck G, Rettberg P, Rabbow E, Strauch W, Seckmeyer G, Facius R,
Reitz G, Strauch K, Schott JU (1996) Biological dosimetry of solar
radiation for different simulated ozone column thickness.
J Photochem Photobiol B Biol 32:189–196