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10.1007@978 3 642 11274 44021 PDF
10.1007@978 3 642 11274 44021 PDF
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
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
have been raised concerning the internal structure of such composition and structure of ice at such extreme pressure
planets, the real existence of a huge ocean and its potential and temperature values. The structure of such an object is
characteristics, and the composition of the atmosphere described in Fig. 1.
allowing its remote identification and study as an exobiology The ocean, with a thickness of about 100 km, lies on
object. At present, only Léger et al. (2004) have published a huge water ice layer (several thousands of kilometers).
a specific study on these objects. As mentioned before, Because of the very high pressure, ice is differentiated.
other studies have been done in a larger context of terres- Because of its intrinsic weight compared to water ice,
trial-type planet structure (that thus includes ocean CO2 is mainly in the form of ice, at the interface between
planets.) In this entry we will consider Léger et al.’s work. the mantle and the water ice layer. This helps prevent CO2
Several supplementary assumptions as limits of their from degassing into the atmosphere, creating to a runaway
model have been done (Léger et al. 2004): greenhouse effect.
● The mass range of planets is limited to 1–8 Earth
Ocean Depth
masses. As biggest objects are easier to detect and
The question of the depth of the ocean was studied con-
characterize, the study focuses on 6–8 Earth mass-
sidering a pure water ocean, whose role is to transfer the
objects.
internal heat of the planet to its surface. Assuming the
● Planets are supposed to form beyond the snowline and
worst case where the temperature gradient is adiabatic
migrate inward to about 1 AU (habitable zone of a
(limit to convection), a surface temperature of the ocean
G star) in about 1 Myr.
of 7 C is obtained. The depth of the ocean is estimated at
● Planets are mostly made of 50% of refractory materials
72 km. This value is higher if the surface temperature of
(Fe, silicates) and 50% of ices according to the com-
the ocean is higher and vice versa. Non-adiabatic depen-
position of protoplanetary nebula (90% of H2O, 5% of
dence of the temperature with depth leads to a smaller
NH3, 5% of CO2). Most of the ice is thus made of
ocean depth.
water.
Atmosphere Composition and Spectrum
Internal Structure Our understanding of the origin of atmospheres (includ-
Modeling of a 6-Earth mass-object has been done starting ing the terrestrial one) is too poor to make accurate pre-
from a model developed to describe the Earth’s interior, dictions of what an ocean planet atmosphere would
taking into account the cooling of the planet, and the exactly contain. However, assuming the composition of
e ∼ 100 km
R/R =
2.00
1.63
e ∼ 20 km
1.24
1.00
0.85
0.69
0.55
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)
Ocean Planet O 1163
cometary material, it is not irrelevant to assume that To determine the density of the planet, one must
volatiles are made of H2O (mainly), NH3, and CO2. In measure its radius and evaluate its mass. The first param-
the atmosphere, NH3 reacts with EUV radiations and eter can be obtained by the transit method. Observation
dissociates leading to H2 and N2. Because of high exo- from space by dedicated satellites (▶ CoRoT, ▶ Kepler,
sphere temperature, H2 escapes and N2 is mixed with the HST, etc) allows the determination of a planetary diameter
remaining atmosphere as a buffer gas. The amount of CO2 at the level of a few percent, assuming simultaneously the
in the atmosphere is strongly limited compared to the knowledge of the stellar diameter with an accuracy of a few
planetary CO2, because most of the CO2 is in the form percent. In principle, the mass of the planet can be deter-
of ice under the water ice layer. Models of ocean planet mined by ▶ radial velocity measurements (RV) based on
atmospheres including photochemical evolution include high spectral resolution spectroscopy. However, the accu-
thus the presence of H2O, CO2, and N2 that should be racy reached by the best RV instruments does not yet allow
detectable in the atmosphere. 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
Exobiological Interest of such Planets
thus very difficult to identify within a large range of
Because of the massive presence of liquid water at their
telluric planet models which one corresponds to the
surface, ocean planets are good candidates for the detec-
observed planet, preventing the observer from identifying
tion of life, assuming the classical criteria for the potential
a real ocean planet (Selsis et al. 2007). Future generations
emergence of life (definition of the habitable zone).
of RV instrument should be more adapted to this quest.
A search for life on these planets requires first an unam-
The knowledge of stellar diameter may also be improved
biguous identification of their nature (see next section),
by interferometric observations.
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
Future Directions
The concept of ocean planet is now considered as
of O2 sinks (such as the Earth’s silicate surface), it appears
a particular case of the bigger family of rocky planets
possible to maintain a quantity of abiotic O2 coming from
(mass between 2 and 10 terrestrial masses), whose
photodissociation of CO2 or H2O in the atmosphere. The
structure and evolution is driven by several parameters
CO2, O2, H2O criteria cannot thus be used. However, in
such as the initial composition, the formation location,
that case, the formation of a large ozone (O3) layer O
and the evolution of orbital parameters. The observational
is limited by perpetual reaction of O3 in the upper
study of ocean planets is thus considered within the
atmosphere with radicals coming from H2O and CO2
framework of detection and characterization of telluric
dissociation. The effective habitability of ocean planets
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.
See also
▶ CoRoT Satellite
▶ Exoplanet, Detection and Characterization
Key Research Findings ▶ Habitable Planet (Characterization)
No observational confirmation of the existence of ocean ▶ Habitable Zone
planets has yet been done. To confirm the existence of ▶ Kepler Mission
these objects, one must determine, with sufficient accu- ▶ Radial Velocity
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-
References and Further Reading
Goldreich P, Tremaine S (1980) Disk-satellite interactions. Astrophys J
eracies should be taken into account carefully. A planet 241:425–441
with a dense core and a hydrogen-rich atmosphere cannot Hayashi C (1981) Structure of the solar nebula, growth and decay of
certainly be distinguished from a H2O-rich planet, but magnetic fields and effects of magnetic and turbulent viscosities on
populations of such objects might be separated by the the nebula. Prog Theor Phys Suppl 70:35–53
Kuchner MJ (2003) Volatile-rich earth mass planets in the habitable zone.
fact that even low-mass hydrogen-rich envelopes produce
Astrophys J 596:L105–L108
very large radii that cannot be attributed to ocean planets. Léger A, Selsis F, Sotin C, Guillot T, Despois D, Mawet D, Ollivier M,
The amount of H2 that can mimic an ocean planet may Labèque A, Valette C, Brachet F, Chazelas B, Lammer H (2004) A new
not be stable, assuming atmospheric escape. family of planets? “Ocean Planets”. Icarus 169:499–504
1164 O Ocean Salinity
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
Ocean, Chemical Evolution of O 1165
atmosphere and oceans and large quantity of Na in the differences with modern seawater, de Ronde et al. (1997)
basaltic proto-crust could have promoted their alkalinity and Channer et al. (1997) noticed that the Cl:Br and Cl:I
by production of Na2CO3, as observed in ▶ soda lakes ratios are higher than in modern seawater and much closer
(Kempe and Degens 1985). However, thermodynamic cal- to the bulk Earth values. This could indicate that Archean
culations indicated that the whole terrestrial available CO2 seawater chemistry was buffered by the mantle through
budget (atmosphere–hydrosphere–mantle) is several emission of dissolved chemical species and volatiles from
orders of magnitude lower than the one needed to make mid-oceanic vents. An alternative hypothesis is the
a global soda ocean (Sleep et al. 2001). absence of planktonic organisms that presently use bro-
Seawater geological record started in the Archean, mine and iodine as metabolites, scavenging them from
though fragmentary and often hampered by large hydro- the ocean.
thermal alteration and metamorphism. The 3.8 Ga ▶ Isua Holland (1984, 2003) used a different approach for
metasedimentary sequences of West Greenland contain constraining the Archean seawater composition, looking
chemical sediments deposited in an oceanic environment at the mineralogy of the deposited sediments. Calcite
without a significant sialic detrital component (Holland (CaCO3) and dolomite (CaMg(CO3)2) were the dominant
2003), i.e., without continental inputs of ionic species. carbonate minerals. ▶ Siderite (FeCO3) was only a
Compelling evidence for the existence of an ocean during common constituent of ▶ BIFs. These observations
deposition of Isua rocks are clastics that have been imply that the Archean ocean was supersaturated with
interpreted as marine turbidites and iron formations respect to CaCO3 and CaMg(CO3)2. For the theoretical
(IFs). These iron-rich layers are magnetite–quartz IFs range of values of atmospheric pCO2 in the Archean, the
and the molar ratio of Fe2O3/FeO is less than 1.0, which pH of seawater was probably between 6 and 7 (Holland
suggests that magnetite was the dominant ▶ iron oxide, 2003; Pinti 2005). The scarcity of siderite in BIFs is also an
and that hematite was absent or very minor in these IFs indicator of a low Fe2þ/Ca2þ ratio in the ocean, i.e., close
(Holland 2003). This in turn suggests that magnetite is not to the ratio of the solubility product of siderite and calcite
a replacement of oxyhydroxides precursors but a primary (4 10–3; Holland 2003). Archean ocean should have been
precipitating phase, thus that the oxygen level in the mainly anoxic as suggested by the mass-independent frac-
oceans was insignificantly low. The oxidation of Fe2þ to tionation (▶ MIF) of the ▶ sulfur isotopes in pre-2.47 Ga
Fe3þ was possibly produced by UV-induced reactions sulfides and sulfates (Farquhar et al. 2000). In the absence
between iron and H2O (Cairns-Smith 1978). The of O2, solar UV interacts with SO2 and generates MIF of
chemistry of Isua seawater is unknown. Appel et al. the sulfur isotopes in the reaction products. In the absence O
(2001) observed ▶ fluid inclusions in quartz globules of atmospheric O2, sulfide minerals would not have been
from pillow breccia at Isua. The chemistry of the highly oxidized during weathering reducing the input of SO2 4 in
saline aqueous fluids (about 25 wt.% NaCl equivalent) the ocean. Another consequence of the low pO2 in the
bears a strong resemblance to present-day seafloor hydro- Archean was the low amount of trace elements such as U,
thermal fluids that likely provoked the alteration of the Mo, and Re that are extremely mobiles in an oxygenated
pillow breccia and the precipitation of quartz. Occur- environment.
rences of fluid inclusions in Archean metasedimentary The Proterozoic ocean was certainly characterized by
rocks see Fluid Inclusions have been signaled in several an increasing O2 concentration, although it reached
terrains of ages spanning from 3.5 to 2.7 Ga (De Ronde appreciable amounts only after 1.8 Ga. The presence of
et al. 1997; Channer et al. 1997; Foriel et al. 2004; O2 certainly favored the scavenging of iron from the ocean
Weiershauser and Spooner 2005). They have been as oxyhydroxides and the deposition of ▶ BIFs (Holland
interpreted as a pristine record of the chemical composi- 1984) and the increasing amount of sulfate. The cessation
tion of Archan seawater. The chemistry of these fluid of BIFs deposition after 1.8 Ga was probably due to
inclusions always pointed out to the mixing between at a decrease in the flux of reductants Fe2þ, H2, and H2S to
least two fluids, a saline fluid of Na–Ca–Cl composition the oceans, which in turn promoted a net increase in the
(seawater) and a hydrothermal effluent, often richer in dissolved O2 content in the deeper ocean (Holland 2006).
Ba and Fe (de Ronde et al. 1997; Foriel et al. 2004). The The end of the Proterozoic was a quite turbulent period
Na:Cl ratio is the same as present-day seawater but NaCl of biological and climatic global changes in the ocean
concentration ranges from 4% to 25% equivalent (against driven by very large glaciations that possibly developed
3.5% equivalent in modern seawater). The salt enrichment in ▶ Snowball Earths.
and the chemistry (Na–Ca–Cl) points out to an evaporate Phanerozoic is the period where the largest (but
seawater residue (Martin et al. 2006). Among the notable discontinuous) geological record of the chemical
1166 O Ocean, Chemical Evolution of
Holland HD, Zimmermann H (2000) The dolomite problem revisited. carbonate or siliceous sediment that disappear close to
Int Geol Rev 42:481–490
the mid-ocean ridges; Layer 2A consisting of 0.5 km thick
Holser WT (1963) Chemistry of brine inclusions in Permian salt from
Hutchinson, Kansas. In: Bersticker AC (ed) Symposium on salt
glassy-to-finely crystalline basalt usually in the form of
(first). Northern Ohio Geological Society, Cleveland, pp 86–95 pillow basalt and 1.5 km thick sheeted vertical diabase
Horita J, Zimmermann H, Holland HD (2002) The chemical evolution of dykes (Layer 2B); Layer 3 (5 km thick and of density
seawater during the Phanerozoic: implications from the record of 3.0 g cm3) made of gabbros (the intrusive form of basalt)
marine evaporites. Geochim Cosmochim Acta 66:3733–3756
and ultramafic cumulates. The crust forms through partial
Kempe S, Degens ET (1985) An early soda ocean? Chem Geol 53:95–108
Knauth LP (2005) Temperature and salinity history of the Precambrian
melting of upwelling mantle beneath mid-ocean spreading
ocean: implications for the course of microbial evolution. centers; the layering results from differentiation of the
Palaeogeogr Palaeoclimatol Palaeoecol 219:53–69 basaltic magma produced by this melting. On Earth,
Martin H, Claeys P, Gargaud M, Pinti D, Selsis F (2006) 6. Environmental the age of oceanic crust ranges from zero at spreading
context. Earth Moon Planet 98:205–245
centers to a maximum of about 180 Ma in the northwestern
McCulloch MT, Bennett VC (1994) Progressive growth of the Earth’s
continental crust and depleted mantle: geochemical constraints.
Pacific.
Geochim Cosmochim Acta 58:4717–4738
Millero FJ (2003) Physicochemical controls on seawater. In: Elderfield H See also
(ed) Treatise on geochemistry, vol 6. Elsevier, Amsterdam, pp 1–21 ▶ Crust
Pinti DL (2005) The formation and evolution of the oceans. In: Gargaud M,
▶ Obduction
Barbier B, Martin H, Reisse J (eds) Lectures in astrobiology. Springer,
Berlin, pp 83–107
▶ Oceans, Origin of
Sleep NH, Zahnle K, Neuhoff PS (2001) Initiation of clement surface ▶ Ophiolite
conditions on the earliest Earth. Proc Nat Acad Sci USA 98:3666–3672 ▶ Plate Tectonics
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 Oceans, Origin of
Cosmochim Acta 53:845–857
Veizer J, Ala D, Azmy K, Bruckschen P, Buhl D, Bruhn F, Carden GAF, DANIELE L. PINTI1, NICHOLAS ARNDT2
Diener A, Ebneth S, Goddéris Y, Jasper T, Korte C, Pawellek F, 1
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l’Atmosphère, Université du Québec à Montréal, O
Weiershauser L, Spooner E (2005) Seafloor hydrothermal fluids, Ben Montréal, QC, Canada
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Nevis area, Abitibi greenstone belt: implications for Archean Maison des Géosciences LGCA, Université Joseph
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Keywords
Comets, cool early Earth, D/H, Isua, Jack Hills zircons,
meteorites, volatiles, water
an ocean of magma. Examples of subsurface oceans are matured during the Precambrian. The presence of d18O of
those predicted for Titan or beneath the smooth icy sur- 7.4 0.7‰ in a 4.4 Ga old detrital zircon of Jack Hills
face of the satellites ▶ Europa and Enceladus. The origin of could thus be the indirect and oldest record of interaction
the ocean entails the complex processes that led to the of a magmatic source with liquid water at the surface of
formation of a stable body of liquid water in thermody- the Earth (Peck et al. 2001). The combination of these
namic equilibrium with an atmosphere on a planetary observations lead to the “▶ cool early Earth” hypothesis
body. The following overview focuses on the formation according to which temperatures were clement, and
of the terrestrial oceans. oceans existed, on the surface of the Earth as early as
4.4 Ga (Valley et al. 2002).
Overview The source of the oceanic water is debated. It is com-
There is good evidence that the oceans were present on the monly accepted that any primitive solar-type atmosphere
▶ Earth in the early ▶ Archean (3.8 Ga ago) after the surrounding the Earth was rapidly blown off by UV and
▶ Late Heavy Bombardment and plausible evidence that X-rays produced by the sun during its proto-(T-Tauri)-
they were already present in the ▶ Hadean. The convinc- star stage. This primitive atmosphere probably did not
ing evidence comes from the two oldest areas of volcanic contain water because Earth accreted too close to the
and sedimentary rocks – the ▶ Isua Supracrustal Belt, Sun. Earth was composed of “dry” raw material related
West Greenland and ▶ Nuvvuagittuq, Quebec, Canada to enstatite chondrite ▶ meteorites (Javoy 1997). The
(formally Porpoise Cove). The ages of the rocks in both present hydrosphere probably was derived from meteor-
areas have been established at about 3.8 Ga (for the latter a itic and cometary material that was added after the moon-
date of 4.3 Ga has been suggested; see ▶ Nuvvuagittuq). In forming impact (Morbidelli et al. 2000). Similarities
the Isua Supracrustal Belt, pillow basalts provide evidence between the hydrogen isotopic ratio D/H of the whole
of underwater eruption, and in both belts, metasedimentary Earth (149 – 153 106), ocean water (155.7 106),
rocks (▶ banded iron formations, metapelite, and ferru- and the average D/H ratios of hydrated carbonaceous
ginous quartzite) are the products of erosion, fluvial trans- chondrites (149 6 106) and Antarctic micrometeor-
port, and subaqueous deposition. Although the setting of ites (154 16 106) indicates that asteroids or asteroi-
some of these rocks may have been lakes or shallow seas, it dal dust are the most likely candidates as water carrier
is far more probable that these rocks erupted or were (Maurette et al. 2000; Marty and Yokochi 2006).
deposited in ocean basins. ▶ Comets apparently exhibit heavier D/H ratios of 290–
The evidence for Hadean oceans is less direct, because 320 106 that are incompatible with oceanic water,
the geological record has been mostly wiped out by the although only a few comets have measured D/H ratios.
intense tectonic activity of the young Earth. Most evidence Mass and isotopic balance calculations suggest that
comes from 4.4 to 4.2 Ga old detrital zircons (ZrSiO4) a maximum of 10–20% of cometary water could have
from the Mt. Narryer quartzite and from ▶ Jack Hills contributed to the terrestrial water budget.
metaconglomerate, Western Australia (Mojzsis et al. The total amount of water delivered to Earth was
2001; Wilde et al. 2001). ▶ Zircon is a common U-rich certainly higher than the mass of the present-day oceans
trace mineral in granitic magmas formed by melting of (1.4 1021 kg), because the losses of volatiles to space
a hydrous source. Parental rocks of the Jack Hills zircons were important at the beginning. Pepin (1991) suggested
possibly formed from melting of subducted hydrated oce- that at least 50% of the water delivered to Earth could have
anic crust. The presence of such a crust requires interac- dissociated to hydrogen and oxygen by the strong early UV
tion between basaltic lava and seawater as is suggested by radiation. Further losses were due to impact erosion of the
the 18O/16O ratio (denoted as d18O) of the Jack Hills atmosphere by arriving planetary bodies. Geophysical and
zircons which extends to 7.4 0.7 ‰. This value can be high-pressure mineralogy experiments suggest that the
explained only by a mixing between the mantle source of modern mantle contains two to five times the ocean vol-
the zircons (d18O ca. 5.0 ‰) and a source enriched in 18O ume of water (Abe et al. 2000) and thus an amount of
compared to the mantle value and derived from water much higher than that preserved in the oceans was
weathering and low-temperature alteration of oceanic added to the newly born Earth.
crust. The d18O values of Hadean, Archaean, and Prote- Theories of how the ocean formed and stabilized on
rozoic zircons show a secular increase to values up to 10‰. the surface of the Earth are based on geophysical modeling
This can be explained by an increase of the amount of and speculation. Water, together with CO2 (now trapped
supracrustal, high-d18O material available for melting and at the Earth’s surface as carbonates), were vapor phases in
assimilation, as the “wet” continental crust evolved and the secondary (outgassed) atmosphere. They did not
Oceans, Origin of O 1169
condense until the surface of the planet cooled down to Future Directions
less than 647 K, the critical point of water. Assuming that Two main questions need to be addressed: (1) When did
all oceanic water was present as a gas, the atmospheric oceans form on Earth? (2) What is the origin of the water?
pressure was 270 bars of H2O plus 40 bars of CO2 (Sleep The first question is difficult to answer because the
et al. 2001; Pinti 2005). This runaway greenhouse atmo- answers must be sought from a period when the geological
sphere, refurbished by a high heat flow from the still record is almost totally absent. Two decades ago the task
molten interior of the Earth, kept the planet’s surface seemed hopeless, but the remarkable results emerging
sufficiently hot for several millions years to prevent water from the Jack Hills zircons and other relicts of the oldest
from condensing on its surface. When a primordial crust crust have radically changed our knowledge of this period.
formed, it reduced the heat flux from the interior and the We need to search for and precisely date and analyze
surface of the Earth cooled down, allowing the condensa- other minerals or rocks that could provide complemen-
tion of water. It is difficult to assess when this occurred. tary evidence of interaction with liquid water. For an
Geophysical models developed by Sleep et al. (2001) sug- answer to the second question, we need to wait for future
gest a few million years or less after the Moon-forming space missions to study and collect samples from the
impact, depending on the parameters chosen. If the geo- surface of comets (e.g., the Rosetta mission) to establish
chemistry of the Jack Hills zircons does indeed reveal the to what extent this material could have contributed to the
presence of liquid water at the surface, this means that terrestrial water budget.
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 See also
survival of the first living communities sometime between ▶ Archea
4.3 Ga and the end of the meteoritic bombardment at ▶ Banded Iron Formation
3.85 Ga. ▶ Comet
The chemistry of seawater in the early Hadean ▶ Cool Early Earth
was likely controlled by high-temperature fluid-rock reac- ▶ Degassing
tions between the hot CO2–H2O-rich atmosphere ▶ Earth, Formation and Early Evolution
and oceans, and primitive basaltic crust. Weathering of ▶ Europa
Na-rich basalt reacting with chlorine saturated the seawa- ▶ Hadean
ter with NaCl. The acidity of water was possibly as high ▶ Isotopic Fractionation (Interstellar Medium) O
as pH = 5 because the ocean was in thermodynamic ▶ Isua Supracrustal Belt
equilibrium with a CO2-rich atmosphere (Pinti 2005), ▶ Jack Hills (Yilgarn, Western Australia)
though alternative models of alkaline oceans saturated ▶ Late Heavy Bombardment
with ▶ thermonatrite formed by weathering of the ▶ Meteorites
basaltic protocrust have been proposed (Kempe and ▶ Nuvvuagittuq (Porpoise Cove) Greenstone Belt
Degens 1985). ▶ Rosetta (spacecraft)
▶ Soda Lakes
Key Research Findings ▶ Thermonatrite
To help understand the origin of the oceans requires the ▶ Titan
integration of information from the Earth, where the ▶ Water, Delivery to Earth
ocean still exists and where a fragmentary record of its ▶ Zircon
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
References and Further Reading
Abe Y, Ohtani E, Okuchi T, Righter K, Drake MJ (2000) Water in the early
shallow oceans that existed there early in the planet’s Earth. In: Righter K, Canup RM (eds) Origin of the earth and moon.
history, and the cold icy satellites provide a record of University of Arizona Press, Tucson, pp 413–433
oceans of very different types. This information can Javoy M (1997) The major volatile elements of the Earth: their origin,
potentially be exported to models for the formation of behavior, and fate. Geophys Res Lett 24:177–180
Kempe S, Degens ET (1985) An early soda ocean? Chem Geol 53:95–108
oceans on exoplanets. Léger et al. (2004), for example, has
Léger A et al (2004) A new family of planets? “Ocean-Planets”. Icarus 169
proposed the existence of a hypothetical family of “Ocean- (2):499–504
Planets” that are entirely covered by an ocean hundreds of Marty B, Yokochi R (2006) Water in the early Earth. Rev Mineral
kilometers deep. Geochem 62:421–450
1170 O Oceanus, Oceani
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
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
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 Oligomer
features labeled.
Definition
The term oligomer is derived from the Greek “oligos,”
See also meaning “a few.” In chemistry, an oligomer is a short
▶ Exoplanets Discovery polymer consisting of approximately five monomer
▶ Microlensing Follow-Up Network units, although agreement as to the strict length cutoff
▶ Microlensing Observations in Astrophysics is debated and varies between four and one hundred.
▶ Microlensing Planets Oligomers are formed by oligomerization, which involves
▶ Optical Gravitational Lensing Experiment linking the monomer units together in a chemical reac-
▶ Probing Lensing Anomalies Network tion. Unlike a ▶ polymer, if one of the repeating units of
oligomer is removed, its chemical properties may be sig-
nificantly altered.
References and Further Reading In biochemistry, the term oligomer is commonly used
Bennett DP et al (2010) Astrophys J 713:837
for short, single-stranded DNA fragments, used in
Gaudi BS et al (2008) Discovery of a Jupiter/Saturn Analog with gravita-
tional microlensing. Science 319(5865):927 ▶ 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
Oligonucleotide O 1173
JACQUES CROVISIER
LESIA - Bâtiment ISO (n 17), Observatoire de Paris, References and Further Reading
Barucci MA, Boehnhardt H, Cruikshank DP, Morbidelli A (2008)
Meudon, France
The solar system beyond Neptune. University of Arizona Press,
Tucson
Dones L, Weissman PR, Levison HF, Duncan MJ (2005) Oort Cloud
Keywords formation and dynamics. In: Festou MC, Keller HU, Weaver H
Comets, small bodies (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
Definition Oort JH (1950) The structure of the cloud of comets surrounding the
The Oort cloud is a spherical cloud of ▶ comets surround- Solar System, and a hypothesis concerning its origin. Bull Astron Inst
ing the Solar System and extending out to heliocentric Neth 11:91–110
distances greater than 100,000 AU. 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,
Overview mŽtŽorites. Bull Astron Ser I 27:194–205
The idea of a distant reservoir of ▶ comets, extending to Van Woerkom AJJ (1948) On the origin of comets. Bull Astron Inst Neth
the verge of the ▶ Solar System, was already discussed by 10:445–472
1176 O Opacity
See also
Opacity ▶ Basalt
▶ Chert
Synonyms ▶ Diagenesis
Attenuation ▶ Hydrothermal Environments
▶ Mars
Definition ▶ Rock
Opacity measures the property of a medium to attenuate ▶ Weathering
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 Oparin’s Conception of
absorption coefficient kn ¼ an =r (with units [cm2 g1]) Origins of Life
which is used.
The main sources of opacity in an astronomical STÉPHANE TIRARD
context are: Faculté des Sciences et des Techniques de Nantes, Centre
● Electron scattering (at temperatures of the order François Viète d’Histoire des Sciences et des Techniques,
1 billion K) EA 1161, Nantes, France
● Electronic transitions (free-free, bound-free, or
bound-bound at temperature of the order 10,000 K)
● Molecular or dust absorption (around or below History
3,000 K) In 1924, Alexander Ivanovitch Oparin (1894–1980),
a young scientist in vegetal physiology, published in Mos-
The opacity plays an essential role in radiative transfer, cow a very important text about the origin of life. With
and thus in the energetic equilibrium of stars. this text, he began a very long career devoted to this
topic. He wrote important books and developed very
See also active experimental researches.
▶ Mean Free Path He published this first text, entitled Origins of Life, in
▶ Optical Depth 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.
Opaline Silica In 1936, Oparin published an important book entitled
Origins of Life. In an interdisciplinary approach, he
Definition suggested a broad scenario of the origins of life on Earth,
Opaline silica (SiO2 · nH2O) occurs at several locations on describing all the steps of the evolution of the Earth, of
▶ Mars, although the definitive identification is conten- matter, and of primordial life. This book knew a great
tious. The confirmed finding would indicate past aqueous success and was translated into several languages, and
activity. Aqueous free silica is a product of ▶ basalt especially into English in 1938.
weathering, when the interaction of water with mafic This book is an exhaustive presentation of Oparin’s
(i.e., Mg- and Fe-rich, silica-poor) ▶ rock rapidly dissolves ideas, very close to his 1924 conception. However, he
olivine, pyroxene, and glass. Opaline silica could have been introduced a lot of current scientific data and two specific
precipitated in hydrothermal or lacustrine (related to points have to be underlined. Firstly, Oparin introduced
a lake) evaporitic environments on Mars. On Earth, it rarely the notion of coacervate coming from Bungenberg de
persists more than a few million years, because it rapidly Jong’s work. Coacervates were microscopical vesicles
transforms into microcrystalline quartz during ▶ diagene- formed in colloidal solutions and Oparin used them as
sis. The persistence of opaline silica on Mars over much a model of a primitive cell in his theory. Secondly, Oparin
longer geologic timescales (billions of years) indicates that claimed that the primitive atmosphere did not contain
water was not present for extended periods of time. CO2, and that it was reductive.
Ophiolite O 1177
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
Orbit O 1179
Definition smaller body will essentially orbit the more massive body.
The orbit of a celestial body is its path around a central This simple mathematical formulation (with modifica-
mass, caused by the acceleration due to the gravitational tions included for general relatively as proposed by Albert
force between them. The planets and minor bodies such as Einstein) forms the backbone of all of celestial mechanics
asteroids and comets in the Solar System orbit the Sun, and dynamical astronomy.
while any celestial body in the Solar System that orbits
something other than the Sun is called a satellite (moon). Overview
Long-term orbital motion is affected by the gravitational As first envisioned by Newton, orbital shapes can be orga-
forces due to all bodies in the vicinity, as well as to torques nized into three different categories based on how much
due to forces such as gas drag, dynamical friction, and energy the orbiting body has: elliptical (including circu-
radiation pressure. lar), parabolic, and hyperbolic. Parabolic and hyperbolic
orbits are unbound, in that they begin and end at an
History infinite distance from the center of mass; the difference
The concept of a celestial orbit can be generalized to between the two is that a body on a parabolic orbit has
include any motion of a celestial body around another zero net energy, while a hyperbolic orbit requires a positive
body, and dates back at least to the earliest written texts. total energy and therefore has a nonzero beginning and
However, Johannes Kepler was the first to establish the ending velocity. Newton realized that these shapes corre-
true qualitative characteristics of astronomical orbits, for- spond to sections of a cone (conic sections), with one
mulating his three laws of planetary motion between 1609 focus of the orbit lying on the rotational axis of the cone.
and 1619. Analyzing the results from painstaking obser- The conic section therefore confines the orbital motion to
vations of the motions of astronomical bodies by Tycho a two-dimensional plane, known as the orbital plane.
Brahe over many years, he determined that these bodies
travel on elliptical orbits (with the Sun at one focus), Basic Methodology
speeding up as they approach the Sun and then slowing It is important to start with a discussion of terminology,
down as they move farther away. As simple as they seem since the language of celestial mechanics (the name for the
today, these two conceptual breakthroughs revolutionized study of orbital dynamics) can become very confusing due
the Western world’s view of the structure of the universe. to the many specialized terms used to describe the prop-
Until this time, the majority view was that the Sun, erties of an orbit.
planets, and star all revolved around the Earth; this theory The general terms for the closest and farthest points of O
is known as geocentrism, with the most well-known pro- an orbit are the periapsis and apoaspsis; these terms can be
ponent being the second-century Greco-Roman philoso- specialized for individual situations (i.e., perigee for orbits
pher Ptolemy. Heliocentrism, the idea that the orbits of around the Earth, perihelion for orbits around the Sun,
the planets are centered on the Sun, had first been pro- and periastron for orbits around any star). The line that
posed by Aristarchos of Samos in the third century BCE, connects the periapsis and the apoapsis is called the line of
but was largely ignored until it was explored by Coperni- apsides; this line naturally passes through the central body,
cus in 1543 and later supported by Galileo’s observations and the direction of the periapsis can rotate to trace
of the motions of Jupiter’s moons. However, these early a circle if the orbit is perturbed (known as apsidal
models conceived the orbits as perfect circles, and there- precession).
fore required many ad hoc additions to fit the observed The motion of an orbiting body can be described by six
motions of planets in the sky. Kepler’s realization that the quantities known as orbital elements. These quantities can
orbits could be elliptical turned heliocentrism into a viable be coordinates of position and velocity, such as the
predictive theory. Cartesian coordinates of a body in a specific reference
Kepler’s third law, that the square of the period of frame; but more commonly orbits are described using
a planet’s orbit is proportional to the cube of the distance the Keplerian orbital elements, since these are independent
from the Sun (P2 / a3), provided the first mathematical of a specific reference direction. The six Keplerian elements
relation between two orbital quantities and formed one of are the semimajor axis and eccentricity of the ellipse, and
the primary drivers for Isaac Newton’s derivation of the the inclination, argument of periapsis, longitude of the
inverse square law of universal ▶ gravitation (Fgrav / 1/r2). ascending node, and either the mean (temporal) or true
Newton was able to show that two gravitationally bound (angular) anomaly (see Fig. 1). The most useful and
bodies will orbit the center of mass of the two-body general elements are semimajor axis, eccentricity, and
system, and if the difference in mass is large enough the inclination, which describe the size, shape, and tilt of the
1180 O Orbit
Celestial body 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
True anomaly to interactions with small planetesimals).
ν
Argument of periapsis
ω
Ω Applications
γ
Longitude of ascending node Reference The study of orbital dynamics is fundamental to much of
direction planetary astronomy in our own Solar System, as well as
Plane of
the study of other stellar and planetary systems. We
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 as they move across the sky over time – the same method
it
Orb has been used since classical times, and we are still using it
to discover ▶ Kuiper Belt objects at the far reaches of the
Orbit. Figure 1 A diagram of an orbit, with several orbital Solar System today. In other planetary systems, we can
elements labeled (inclination, argument of periapsis, measure the radial velocity of the central star and use
longitude of the ascending node, and true anomaly). The that motion to calculate the orbital motion of surround-
additional orbital elements not labeled are the semimajor axis ing planets even when they are impossible to detect
(i.e., size of the orbit) and the eccentricity (i.e., ellipticity) of the directly.
orbit (WikiCommons) We can also explore the evolution of the orbits of
bodies in planetary systems over time using computer
simulations. Computational numerical simulations of
orbit with respect to the reference orbital plane (the plane planetary dynamics all aim to solve the same general
of reference in the Solar System is known as the ecliptic problem: how to calculate the gravitational and nongravi-
plane). The semimajor axis is defined as one half of the tational forces on a system of bodies accurately over long
major axis of the orbital ellipse; note that this is different timescales (known as the n-body problem). The solution
than the distance to the central body when the orbit is not to the dynamical evolution of any system of three or more
a circle. The eccentricity describes how elliptical the orbit bodies cannot be solved exactly, but there are many dif-
is, ranging from 0 for a circular orbit to 1 for a parabolic ferent types of mathematical algorithms that have been
orbit. The inclination of the orbit is measured in degrees, developed to optimize the pursuit of an approximate
ranging from 0 to 180 (values larger than 90 indicate a solution (i.e., Runge–Kutta, symplectic integration, etc).
“retrograde” orbit that moves in the opposite direction Important applications for these tools include the study of
around the star). The other elements are less intuitive and the formation of planetary systems through long-term
less frequently used, since they primarily define a single accretion of planetary embryos, as well as the dynamical
orbital position and average out to zero for a population of history of populations of small bodies in our Solar System
randomly oriented bodies (as opposed to eccentricity and (such as the Kuiper Belt).
inclination that have a characteristic distribution for
a randomized population), and they will not be discussed See also
in detail here. ▶ Apsidal Angle
There are a number of processes that can lead to an ▶ Gravitation
evolution of the orbital parameters of a single body or ▶ Kuiper Belt
a population of bodies; these effects are known as pertur- ▶ Orbit
bations. These perturbations generally fall into two cate- ▶ Period
gories: gravitational perturbations due to interactions with ▶ Planet Formation
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 References and Further Reading
drag or radiation pressure). Gravitational forces between Danby JMA (1992) Fundamentals of celestial mechanics. Willmann-Bell
Organelle O 1181
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 also implied an anticlerical political stance (Farley 1977;
two worlds, and the appearance of life on Earth may Fry 2002; Strick 2009).
therefore be better envisioned as a continuum that Pasteur’s results made it difficult to explain the origin
seamlessly joins the prebiotic synthesis and accumulation of life by means of spontaneous generation, a conclusion
of organic molecules in the primitive environment with that was interpreted as a major blow to one of the implicit
the emergence of self-sustaining, replicative chemical sys- assumptions underlying Darwin’s views. Several devoted
tems capable of undergoing Darwinian evolution. 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
Overview Helmholtz, felt that they could sidestep the issue by
The term “origin of life” has several possible meanings, but
assuming that viable microbes had been delivered to the
from the perspective of contemporary biology and within
primitive Earth by meteorites, attempting to keep the
the framework of evolutionary theory it refers to the first
validity of evolution. In fact, toward the end of the nine-
appearance of primordial living entities. It is generally
teenth century, the belief that life on Earth had evolved
assumed that since antiquity early philosophers and nat-
from extraterrestrial organisms elicited a number of pro-
uralists appealed to spontaneous generation to explain the
posed mechanisms that could have transported microbes
origin of life. However, with few exceptions, this was not
between planets (see, e.g., Arrhenius 1908), but little
the case. Spontaneous generation was seen mostly as
attention was given to the central issue of their actual
a nonsexual reproductive mechanism, and it was not
origin (Oparin 1924, 1938; Kamminga 1982; Fry 2002).
until Erasmus Darwin, Georges Louis Leclerc de Buffon,
The recent revival of the panspermia hypotheses has
and Jean-Baptiste de Lamarck incorporated spontaneous
not changed this situation. Recent proposals that terres-
generation within their transformist views that it was
trial life arose elsewhere and was transferred to our planet
conceived as the mechanism that had led to the first
are based on the well-established exchange of solid mate-
appearance of life on our planet (Farley 1977).
rial among the different bodies in the solar system, the
Like his predecessors, Charles Darwin surmised that
assumption of more benign environments in some of
plants and animals arose naturally from primordial
these other planets during the early stages of planetary
nonliving matter, but like many others he rejected the
system, and the surprising resistance of a number of pro-
idea that putrefaction of preexisting organic compounds
karyotic species to environmental insults such as high
could lead to the appearance of organisms. Darwin’s let-
fluxes of UV light or corpuscular radiation. There are of
ters, notebooks, and few published statements on the issue
course alternative explanations for these observations, but
demonstrate that although he favored the possibility that
with formidable disregard for plausibility, the panspermia
life could appear by natural processes from simple inor-
hypothesis has been recently restated in a variety of con-
ganic compounds, his reluctance to discuss it in public was
texts. However, its current advocates, like those of the late
due at least in part to the recognition that it was difficult to
nineteenth century, simply transfer the question of the
develop fruitful experimental approaches to address the
origin of life to another habitable planet in our galaxy
problem. Nevertheless, his theory established the frame-
without providing an explanation of how it happened.
work that would lead to a number of attempts to explain
the origin of life by introducing principles of historical
The Origin of Life: A Question Without
explanation (Peretó et al. 2009).
an Answer?
A century and a half after Darwin admitted how little was
Panspermia: Past and Present understood about the origin of life, we still do not know
Darwin’s Origin of Species was published in 1859, the very when and how the first living beings appeared on Earth.
same year in which Pasteur began the experiments that Despite the seemingly insurmountable obstacles sur-
would lead him to disprove spontaneous generation of rounding the understanding of the origin of life, or per-
microbes. His results had implications that went well haps because of them, there has been no shortage of
beyond the limits of academia. Since the times of Lamarck discussion about how it took place. Since the attributes
and Buffon spontaneous generation had been associated of the first living entities are unknown, it is not surprising
in France not only with evolutionary theory but also with that an inventory of current views on the origin of life
secular attitudes and radical political views. In such an reveals a mixture of opposites of every kind.
entangled atmosphere, spontaneous generation embodied Looking back across time since the origin of life
not only support for evolution and transformist views, but occurred is a scientific exercise fraught with guesswork,
Origin of Life O 1185
since the evidence required to describe the prebiotic envi- translation apparatus, including some late steps in the
ronment and the nature of the events that led to the first development of the ▶ genetic code. Such comparative
living systems is scant and difficult to understand. Since approaches have made it possible to distinguish the ori-
most of the rocks from early Archean times that have been gin-of-life problem from a whole series of other issues,
preserved have been metamorphosed to a considerable often confused, that belong to the domain of the early
extent, there is no direct evidence of the environmental evolution of microbial life.
conditions on the Earth at the time of the emergence of Molecular phylogenies, comparative genomics, and
life, nor any fossil register of the predecessors of the first bioinformatics approaches fail to provide direct informa-
cells. There is no direct information of the chemical com- tion on the nature of the first living beings. A cladistic
position of the terrestrial atmosphere during the period of approach to the origin of life is not feasible, since all
the origin of life, nor of other general and local environ- possible intermediates that may have once existed have
mental conditions, which may have been important for long since vanished. Phylogenetic analyses based on com-
the appearance of living systems (Bada and Lazcano 2009). parative genomics provide important clues to early stages
Moreover, any explanation of the origin of living sys- of biological evolution, but at the time being their appli-
tems should attempt, at least implicitly, the definition of cability cannot be extended beyond a threshold that cor-
a set of minimal criteria for what constitutes a living responds to a period of cellular evolution in which protein
organism. However, this has proven to be an elusive intel- biosynthesis was already in operation, that is, the RNA/
lectual endeavor, though not for lack of trying. The protein world (Becerra et al. 2007).
absence of such a definition sometimes gives the impres- Although there have been attempts to deduce the
sion that what is meant by the origin of life is described in nature of the first living systems from extant metabolism
somewhat imprecise terms, and that several entirely dif- (Wächtershäuser 1988), there is a lack of simple continuity
ferent and even opposing questions are often confused between the biosynthetic and the (possible) prebiotic
(Lazcano 2008). Prior to the discovery of catalytic activity pathways (Lazcano and Miller 1999). For instance, it is
of RNA molecules, for instance, the origin of the genetic likely that the Strecker synthesis and the Bucherer–Bergs
code and of protein synthesis was considered synonymous reaction played a major role in the prebiotic accumulation
with the appearance of life itself. This is no longer of amino acids, but these abiotic routes are very different
a dominant point of view: four of the central reactions from transamination and the reverse Krebs cycle found in
involved in protein biosynthesis are catalyzed by extant cells. The prebiotic synthesis of purines is from
ribozymes, and their complementary nature suggests HCN and its derivatives, and not from glycine, formate, O
that they first appeared in an ▶ RNA world (Yarus 2010), and NH3, as in current biology. It is true that the amino
that is, that ribosome-catalyzed, nucleic acid-coded pro- imidazole carboxamide ribotide in the de novo purine
tein synthesis is the outcome of Darwinian selection of nucleotide biosynthetic pathway is comparable to the
RNA-based biological systems, and not of mere physico- amino imidazole carbonitrile formed in the prebiotic
chemical interactions that took place in the prebiotic pathway, and a few other examples are found in the for-
environment. mation of uracil, glutamic acid, pyrroles, pyrimidines, and
acetic acids (Lazcano 2010a). However, the similarities
Phylogenomics and Extant Metabolism Do between these abiotic reactions and their enzyme-
Not Explain the Origin of Life mediated counterparts in biosynthetic pathways do not
The variations of traits common to extant species can be necessarily indicate an evolutionary continuity between
explained as the outcome of divergent processes from prebiotic chemistry and biochemical pathways, but may
ancestral life forms that existed prior to the separation of simply reflect chemical determinism. In other words, the
the three major biological domains, that is, the last com- few cases in which some extant metabolic pathways and
mon ancestor of Bacteria, Archaea, and Eucarya. This prebiotic chemical processes share similar steps may be
entity was likely far removed, if not in geological time, at due to the unique way in which given reactions can
least in complexity from the first living systems. Although take place.
no evolutionarily intermediate stages or ancient simplified
versions of the basic biological processes have been dis- From the Primitive Soup to the RNA World
covered in contemporary organisms, the differences in the It is generally believed that after Louis Pasteur had
structure and mechanisms of gene expression and replica- disproved the spontaneous generation of microbes using
tion among Bacteria, Archaea, and Eucarya have provided his famous swan-necked flask experiments, the discussion
insights into the stepwise evolution of the replication and of life’s beginning had been banished to the realm of
1186 O Origin of Life
useless speculation. However, scientific literature from the published in 1938 (Oparin 1938). In this book, Oparin’s
first part of the twentieth century shows the many original proposal was revised, leading to the assumption of a
attempts by major scientists to solve this problem. The highly reducing milieu in which iron carbides of geological
list covers a rather wide range of explanations that go from origin would react with steam to form hydrocarbons. Their
the ideas of Pflüger on the role of hydrogen cyanide on the oxidation would yield alcohols, ketones, aldehydes, etc., that
origin of life, to those of Svante Arrhenius on panspermia, would then react with ammonia to form amines, amides,
and includes Leonard Troland’s hypothesis of a primordial and ammonium salts. The resulting protein-like com-
enzyme formed by chance events in the primitive ocean, pounds and other molecules would form a hot dilute
Alfonso L. Herrera’s sulfocyanic theory on the origin of soup, in which they would aggregate to form colloidal
cells, Harvey’s 1924 suggestion of an heterotrophic origin systems such as coacervates, from which the first heterotro-
in a high-temperature environment, and the provocative phic microbes evolved. Like many others at the time, Oparin
1926 paper that Hermann J. Muller wrote on the abrupt, did not address in his 1938 book the origin of nucleic acids,
random formation of a single, mutable gene endowed because their role in genetic processes was not even
with catalytic and autoreplicative properties (cf. Lazcano suspected. Because of this, inheritance of primordial genetic
1992; 2010b). information was assumed by Oparin to be the result of
In spite of their diversity, most of these explanations growth and division in the coacervate drops he advocated
went unnoticed, in part because they were incomplete, as models of precellular systems (Lazcano 2010b).
speculative schemes largely devoid of direct evidence and Similar ideas were being developed independently by
not subject to fruitful experimental testing. Although other researchers. In 1929, John B. S. Haldane, the extraor-
some of these hypotheses considered life as an emergent dinary British polymath, also argued that the origin of life
feature of nature and attempted to understand its origin had been preceded by the synthesis of organic compounds.
by introducing principles of historical explanation, the Based on experiments by E. C. C. Baly, an English chemist
dominant view was that the first forms of life had been who claimed that he achieved the syntheses of amino acids
photosynthetic microbes endowed with the ability to fix and sugars by the UV irradiation of a solution of CO2 in
atmospheric CO2 and to use it with water to synthesize water, Haldane suggested that the absence of oxygen in
organic compounds. A major scientific breakthrough a CO2-rich primitive atmosphere had led to the synthesis
occurred, however, when Oparin (1924) suggested of organic compounds and the formation of a “hot dilute
a heterotrophic origin of life that assumed that prior to soup.” Haldane was also influenced by D’Herelle’s discov-
the emergence of the first cells a prebiotic synthesis of ery of phages, and suggested that viruses represented an
organic compounds led to the accumulation of the prim- intermediate step in the transition from the prebiotic soup
itive broth. to the first heterotrophic cells. Life may have remained,
Such ideas were supported not only by the evidence of wrote Haldane (1929), in the viral stage for many millions
organic compounds in meteorites, but also by the striking of years before leading to the first cells.
nineteenth-century experimental demonstrations that The hypothesis that the first organisms were anaerobic
biochemical compounds such as urea, alanine, and sugars heterotrophs is based on the assumption that abiotic
could be formed under laboratory conditions, as had been organic compounds were a necessary precursor for the
demonstrated by Wöhler, Strecker, and Butlerow, respec- appearance of life. Experimental evidence in support of
tively. Oparin’s proposal, which was based on his Darwin- Oparin’s proposal of chemical evolution came first from
ian credence in a gradual, slow evolution from the simple Harold C. Urey’s laboratory, whom had been involved
to the complex, stood in sharp contrast with the then with the study of the origin of the solar system and the
prevalent idea of an autotrophic origin of life. Since chemical events associated with this process. Urey had also
a heterotrophic anaerobe is metabolically simpler than considered the origin of life in the context of his proposal
an autotrophic one, the former would necessarily have of a highly reducing terrestrial atmosphere (Urey 1952).
evolved first. Thus, based on the simplicity and ubiquity The first successful prebiotic amino acid synthesis was
of fermentative reactions, Oparin (1924) suggested in carried out with an electric discharge and a strongly reduc-
a small booklet that the first organisms must have been ing model atmosphere of CH4, NH3, H2O, and H2 (Miller
heterotrophic bacteria that could not make their own food 1953). The result of this experiment was a large yield of
but rather consumed organic material present in the prim- a racemic mixture of amino acids, together with hydroxy
itive milieu. acids, short aliphatic acids, and urea. One of the surprising
These ideas were further elaborated and refined in results of this experiment was that the products were not
a more extensive book whose English translation was a random mixture of organic compounds; rather,
Origin of Life O 1187
a relatively small number of compounds were produced in the years that followed the ▶ Miller–Urey experiment
substantial yield. Moreover, with a few exceptions, the attempts to understand the origin of life were shaped to
compounds were of biochemical significance. a considerable extent by the unraveling of the molecular
It is very unlikely, however, that the RNA world would details of DNA replication and protein biosynthesis.
have arisen from such abiotic organic syntheses. The dis- Perhaps not surprisingly, Herman J. Muller profited
covery of catalytically active RNA molecules gave consid- from the rapid developments in molecular biology and
erable credibility to prior suggestions that the first living the success in prebiotic syntheses to update his gene-first
organisms were largely based on ribozymes, a hypothetical proposal by arguing that what had emerged in the prim-
stage called the RNA world (Gilbert 1986; Joyce 2002). itive oceans had been, in fact, a primordial DNA. For
This possibility is now widely accepted, but the problems Muller and his followers, the essence of life lies in the
involved with the synthesis and accumulation of RNA combination of autocatalysis, heterocatalysis, and muta-
components suggest that ribozymes were not a direct out- bility, which he equated with evolutionary potential. In
come of prebiotic evolution, but may have been one of the doing so, he implicitly stated that life could be so well
evolutionary outcomes of what are now referred to as pre- defined that the exact point at which it started could be
RNA worlds. However, the chemical nature of the first established with the sudden appearance of the first DNA
genetic polymers and the catalytic agents that may have molecule (Muller 1961).
formed the pre-RNA worlds that bridged the gap between On the other hand, Wächtershäuser (1988) has argued
the prebiotic broth and the RNA world are completely that life started with the appearance of an autochemo-
unknown and can only be surmised. lithotrophic two-dimensional monomolecular layer bound
The catalytic versatility of RNA molecules clearly to pyrite that lacked a genetic system. According to this
merits a critical reappraisal of Muller’s viewpoints (Muller proposal, the synthesis in activated form of organic com-
1961). However, the discovery of ribozymes does not pounds such as amino acid derivatives, thioesters, and
imply that autocatalytic nucleic acid molecules ready to keto acids is assumed to have taken place on the surface
be used as primordial genes were floating in the primitive of FeS and FeS2 in environments that resemble those of
oceans, or that the RNA world emerged completely assem- deep-sea hydrothermal vents. Wächtershäuser’s explicit
bled from simple precursors present in the prebiotic soup. adherence to Karl Popper’s philosophical stance (Popper
The evidence supporting the presence of a wide range of 1959) played a major role in shaping his ideas. The
organic molecules on the primitive Earth, including mem- hypothetico-deductive system developed by Popper under-
brane-forming compounds, suggests that the evolution of lines at least in part Wächtershäuser’s so-called methodol- O
membrane-bounded molecular systems preceded cellular ogy of retrodiction, which he has used to argue that “lines
life on our planet, and that life is the evolutionary out- of descent can be traced back to the first archaic pathways.”
come of a process, not of a single, fortuitous event. According to this scheme, chemoautotrophic carbon fixa-
tion took place by a reductive citric acid cycle, or reverse
A Modern Scientific Battlefield Krebs cycle, of the type originally described for the photo-
As summarized by Eschenmoser (2008), two major camps synthetic green sulfur bacterium Chlorobium limicola.
can be recognized among those working on the origins of Wächtershäuser’s insightful prediction that ferrous
life, that is, those assuming that the emergence of auto- sulfide in the presence of hydrogen sulfide (H2S) is an
catalytic “metabolic” cycles in the primitive Earth was efficient reducing agent should not be underappreciated.
essential for the appearance of genetic systems, and those Pyrite formation can produce molecular hydrogen, pro-
that assume the priority of genetic polymers endowed mote the formation of ammonia from dinitrogen, and can
with catalytic properties. These two different viewpoints reduce a few organic molecules under relatively mild
reflect a rather sharp division between those who favor the conditions. However, compared with the surprising vari-
idea that life is an emergent interactive system endowed ety of biochemical compounds that are readily synthesized
with dynamic properties that exist in a state close to in one-pot Miller–Urey type simulations, the suite of
chaotic behavior, and those who are reluctant to adhere molecules produced under the conditions suggested by
to a definition of living systems lacking of a genetic com- Wächtershäuser is quite limited. Wächtershäuser’s proposal,
ponent whose properties reflect the role that Darwinian which is part of a trend that assumes that genetic material
natural selection and, in general, evolutionary processes, was not essential for the origin of life, has been modified by
have played in shaping its central characteristics. Russell and Hall (1997) and Martin and Russell (2003),
With few exceptions, such as the views advocated by who have argued that a primitive version of the acetyl-
Sidney W. Fox and others (cf. Fox and Dose 1977), during CoA pathway existed within FeS-rich mineral boundaries.
1188 O Origin of Life
Based on the hypothesis that core metabolic processes self-organizing metabolic cycles that at first did not
have not changed since the emergence of life, Morowitz require genetic polymers.
(1992) has also argued that intermediary metabolism The available evidence suggests that the origin of life
recapitulates prebiotic chemistry. He maintains that the and the onset of natural selection resulted from self-
basic traits of metabolism could only evolve after the organization phenomena involving nonequilibrium
closure of an amphiphilic bilayer membrane into physical and chemical processes that go beyond simple
a vesicle, that is, that the appearance of membranes rep- Newtonian physics, and that were based on the abiotic
resents the discrete transition from nonlife to life. synthesis, stability, and accumulation of organic mole-
According to his hypothesis, reverse Krebs cycle- cules endowed with differential reactivity, on nucleic
dependent life appeared with “minimal protocells” acid self-assembly and chemistry, and the spontaneous
formed by bilayer vesicles made up of small amphiphiles assembly of prebiotic amphiphiles into micelles and
and endowed with pigments capable of absorbing radiant bilayer membranes, among others (de Duve 2005; Lehn
energy stored as a chemiosmotic proton gradient across 2002; Budin and Szostak 2010). The many examples of
the membrane. self-organizing physical systems that lead to highly
ordered structures suggest that, in addition to natural
Life as an Emergent Complex System selection, there are other mechanisms of ordered complex-
The development of metabolic approaches to the origin of ity that may have played a role in the origin of living
life like that of Morowitz (1992) and his followers is systems. The recognition that the emergence of life is the
increasingly based on complexity theories and self- outcome of an evolutionary process constrained by the
assembly phenomena. The background of current meta- laws of physics and chemistry can lead to the acceptance
bolic views lies not in Oparin’s proposals, but in the that many properties associated with living systems, such
attempt to extrapolate to biology the deeply rooted ten- as replication, self-assembly, or nonenzymatic catalysis are
dency in physical sciences to search for all-encompassing also found in nonliving entities (Lazcano 2010a, b). More-
laws that can be part of grand theory that can explain over, if the origin of life is seen as the evolutionary tran-
many, if not all, complex systems (Lazcano 2010a). These sition between the nonliving and the living, then it is
explanations of the origin of life are based on the idea meaningless to attempt to draw a strict line between
that the emergence of autocatalytic “metabolic” cycles these two worlds, and the appearance of life on Earth
on the primitive Earth was an essential prerequisite for should, therefore, be seen as an evolutionary continuum
the appearance of genetic systems. According to this that seamlessly joins the prebiotic synthesis and accumu-
approach, life can be considered an emergent interactive lation of organic molecules in the primitive environment,
system endowed with dynamic properties that exist in with the emergence of self-sustaining, replicative chemical
a state close to chaotic behavior. Some of these proposals systems capable of undergoing Darwinian evolution
reflect a reaction against the reductionism of molecular (Lazcano 2010a, b).
biology, as well as the adherence to all-encompassing
views based on complexity theories and self-assembly Conclusions
that tend to be more popular among physical scientists The remarkable coincidence between the monomeric con-
than among biologists. stituents of living organisms and those synthesized in
Of course, the many examples of self-organizing phys- laboratory simulations of the prebiotic environment
ical systems that lead to highly ordered structures demon- appears to be too striking to be fortuitous. Nevertheless,
strate that, in addition to natural selection, there are other at the time being the gap between the primitive soup and
mechanisms of ordered complexity that operate. Self- the RNA world is discouragingly large. Life cannot be
assembly is not unique to biology, and may indeed be reduced to one single molecule such as DNA or a popula-
found in a wide variety of systems, including cellular tion of replicating ribozymes, but present-day biology
automata, the complex flow patterns of many different indicates that it could not have evolved in the absence of
fluids, in cyclic chemical phenomena (such as the a genetic replicating mechanism ensuring the stability and
Belousov–Zhabotinsky reaction) and, quite significantly, diversification of its basic components.
in the autoorganization of lipidic molecules in bilayers, As shown by the work of Jack Szostak and his associ-
micelles, and liposomes. There are indeed some common ates (Mansy et al. 2008), it is possible to overcome at least
features among these systems, and it has been suggested in part the intellectual dichotomy between those claiming
that “emergent properties” or “self-organizing principles” that the appearance of the first life forms depended
suffice to explain the origin of life with the emergence of on informational oligomeric compounds, that is, the
Origin of Life O 1189
so-called genetic approach, and those that argue that it Budin I, Szostak JW (2010) Expanding roles for diverse physical phenom-
ena during the origin of life. Annu Rev Biophys 39:245–263
was based on autocatalytic metabolic cycles. Instead of
De Duve C (2005) The onset of selection. Nature 433:581–582
engaging in footling arguments about when exactly did Eschenmoser A (2008) On a hypothetical generational relationship
life start, the recognition that it is the outcome of an between HCN and constituents of the reductive citric acid cyle. In:
evolutionary process constrained by the laws of physics Herdewijn P, Kisakürek V (eds) Origin of life: chemical approach.
and chemistry can lead to the acceptance that many prop- Wiley-VCH, Zürich, pp 31–50
Farley J (1977) the spontaneous generation controversy from descartes to
erties associated with living systems, such as replication,
oparin. John Hopkins University Press, Baltimore/London
self-assembly, or catalysis are also found in nonliving Fox SW, Dose K (1977) Molecular evolution and the origin of life. Marcel
entities. Dekker, New York
Perhaps we will never know exactly how life origi- Fry I (2002) The emergence of life on earth. Rutgers University Press,
nated. As in other areas of evolutionary biology, answers New Brunswick
Gilbert W (1986) The RNA world. Nature 319:618
to questions on the origin and nature of the first life forms
Haldane JBS (1929) The origin of life. Rationalist Annu 148:3–10
can only be regarded as inquiring and explanatory rather Joyce GF (2002) The antiquity of RNA-based evolution. Nature 418:
than definitive and conclusive. This does not imply that all 214–221
origin-of-life theories and explanations can be dismissed Kamminga H (1982) Life from space – a history of panspermia. Vistas
as pure speculation, but rather that the issue should be Astron 26:67–86
Lazcano A (1992) Origins of life: the historical development of recent
addressed conjecturally, in an attempt to construct not
theories. In: Margulis L, Olendzenski L (eds) Environmental evolu-
a mere chronology but a coherent historical narrative by tion: effects of the origin and evolution of life on planet earth. MIT
weaving together a large number of observations and Press, Cambridge, pp 57–69
experimental results (Kamminga 1982). 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
See also 40:161–167
▶ Baly’s Experiment Lazcano A (2010b) Historical development of origins of life. In: Deamer
▶ Bernal’s Conception of Origins of Life DW, Szostak J (eds) Cold spring harbor perspectives in biology:
▶ Calvin’s Conception of Origins of Life the origins of life. Cold Spring Harbor Press, Cold Spring Harbor,
▶ Cell pp 1–16
Lazcano A, Miller SL (1999) On the origin of metabolic pathways. J Mol
▶ Chemical Evolution
Evol 49:424–431
▶ Complexity Lehn JM (2002) Towards self-organization and complex matter. Science
▶ Darwin’s Conception of Origins of Life 295:2400–2403
O
▶ Evolution (Biological) Mansy SS, Schrum JP, Krishnamurthy M, Tobé S, Treco DA, Szostak JW
▶ Genetics (2008) Template-directed synthesis of a genetic polymer in a model
protocell. Nature 454:122–125
▶ Life
Martin W, Russell MJ (2003) On the origin of cells: a hypothesis for the
▶ Metabolism (Biological) evolutionary transition s from abiotic chemistry to chemoautotro-
▶ Monod’s Conception on the Origins of Life phic prokaryotes, and from prokaryotes to nucleated cells. Philos
▶ Oparin’s Conception of Origins of Life Trans R Soc Lond B Biol Sci 358:59–85
▶ Panspermia Miller SL (1953) A production of amino acids under possible primitive
Earth conditions. Science 117:528
▶ Phylogeny
Morowitz HJ (1992) Beginnings of cellular life: metabolism recapitulates
▶ Prebiotic Chemistry biogenesis. Yale University Press, New Haven
▶ Replication (Genetics) Muller HJ (1961) Genetic nucleic acid: key material in the origin of life.
▶ RNA World Persp Biol Med 5:1–23
▶ Urey’s Conception of Origins of Life Oparin AI (1924) Proiskhozhedenie Zhizni. Mosckovskii Rabochii,
Moscow. Reprinted and translated in Bernal JD (1967) The origin
of life. Weidenfeld and Nicolson, London
References and Further Reading Oparin AI (1938) The origin of life. McMillan, New York
Arrhenius S (1908) Worlds in the making: the evolution of the Universe. Peretó J, Bada JL, Lazcano A (2009) Charles Darwin and the origins of life.
Harper & Row, New York Orig Life Evol Biospheres 39:395–406
Bada JL, Lazcano A (2009) The origin of life. In: Ruse M, Travis J (eds) The Popper K (1959) The logic of scientific discovery. Basic Books,
harvard companion of evolution. Belknap/Harvard University Press, New York
Cambridge, pp 49–79 Russell MJ, Hall AJ (1997) The emergence of life from iron monosulphide
Becerra A, Delaye L, Islas A, Lazcano A (2007) Very early stages of bubbles at a submarine hydrothermal redox and pH front. J Geol Soc
biological evolution related to the nature of the last common ances- Lond 154:377–402
tor of the three major cell domains. Annu Rev Ecol Evol Syst Strick JE (2009) Darwin and the origin of life: public versus private
38:361–379 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
Orpheus
Ornithine ▶ Theia
Synonyms
Diaminovaleric acid
Ortholog
Definition
L-Ornithine is a natural, biological, non-coded diamino ▶ Orthologous Gene
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 Orthologous Gene
cycle. Ornithine can be derived from glutamate and is used
in the biosynthesis of arginine and is therefore indirectly Synonyms
coupled to protein synthesis in mammals. Polypeptides Ortholog
containing unprotected ornithines undergo spontaneous
lactamization. L-ornithine is produced through hydrolysis Definition
of citrulline and L-arginine in alkaline medium, that is, the Orthologous genes (or orthologs) are a particular class of
reverse of the biosynthetic process or the same direction homologous ▶ genes. They are found in different species
observed in the urea cycle. Both, L- and D-ornithine have and have diverged following the speciation of the species
been tentatively identified in the Murchison meteorites hosting them. Therefore, orthologous genes in different
(Meierhenrich 2004), (concentration < 5ppb). species derive from a ▶ common ancestral gene found in
Molecular formula: C5H12N2O2 the ancestor of those species. Given their common origin,
Molecular mass: 132.2 g·mol1 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,
O
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
H2N OH
using molecular phylogeny.
NH2
Ornithine
See also
▶ Common Ancestor
Ornithine. Figure 1 ▶ 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
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
See also
Oxirane ▶ Dioxygen
▶ Oxidation
▶ Ethylene Oxide ▶ Oxygen Isotopes
1196 O Oxygen (Molecule)
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 at the inner edge of the solar nebula, in the proximity of the
isotopically separated absorption bands leading to its pho- Sun. Later models assume that the CO self-shielding
todissociation through the predissociation mechanism. In occurred at the cold environment, at the envelope of the
a gas medium irradiated by an ultraviolet light, the wave- outer solar nebula (Lyons and Young 2005) or in the paren-
lengths which correspond to the absorption bands for tal molecular cloud, part of which later evolved to the solar
12 16
C O, the dominantly major isotopologue of CO, are nebula (Yurimoto and Kuramoto 2004). In these CO self-
more quickly attenuated than those for other minor shielding models, the 17,18O-rich isotope signature pro-
isotopologues. The CO self-shielding is considered to be duced by the self-shielding is assumed to be transmitted to
responsible for the carbon and oxygen isotope fraction- H2O, and finally trapped by the silicate minerals.
ations in molecular clouds (Visser et al. 2009). Many We may test these O isotope fractionation models
workers attempted to apply the self-shielding effect to by the knowledge of several important end-member
the early solar nebula. First, Kitamura and Shimizu components. The most important one is the solar com-
(1983) proposed that 17,18O-rich O atoms derived from position. The solar composition here refers to the compo-
the self-shielding effect in the photodissociation of O2, sition of the solar gas at the surface of the Sun. The
exchanged with mineral grains upon shock heating, may temperature at the solar surface is low enough to prevent
lead to 17,18O-enriched planetary solid materials. How- occurrence of any nuclear reactions to alter the O-isotope
ever, Navon and Wasserburg (1985) pointed out that the composition. Also, the mixing of masses between the
17,18
O-enrichment produced by the self-shielding will be surface and the solar core is considered to be negligibly
rapidly erased by the reaction between the O atom and O2. small. Therefore, the composition of the gas at the
Clayton (2002) proposed that CO self-shielding may occur solar surface is considered to preserve the proto-solar
Oxygen Isotopes O 1199
composition, representing the bulk composition of the contains pieces of organic particles that display striking
solar nebula. The MIF models assume that the solar enrichments in the 17O and 18O at almost paralleled
O-isotope composition is the same with the terrestrial degrees, by as high as 530‰ relative to the terrestrial
composition, whereas the CO self-shielding models composition. The oxygen isotope anomalies were associ-
assume a 16O-rich solar composition. Hashizume and ated with carbon isotope anomalies, roughly with
Chaussidon (2005) discovered at the surface of several a relationship of d17O 1.05 d18O 0.5 d13C.
metallic grains contained in lunar soil samples, irradiated By the CO self-shielding, the enrichment of 13C is
by the solar wind at the lunar surface since 1–2 Gyr ago, an indeed expected to be associated with the 17,18O enrich-
oxygen component enriched in 16O by >2% relative to ment. However, the 13C-rich signature produced among
the terrestrial composition. They assigned the observed the photodissociated products is considered to be
16
O-rich oxygen to the solar-wind composition, conclud- compromised by a competing reaction. The ion molecule
ing that the bulk solar nebula was 16O-rich. However, the reaction (12CO + 13C+ ↔ 12C+ + 13CO + 35 K) is partic-
situation was complicated by the finding of a distinct ularly active at low temperatures, whose isotope effect is
O-isotope composition at the surface of metallic grains considered to exceed the effect by the self-shielding below
from a different lunar soil sample, where O was enriched a gas temperature of 60 K (Visser et al. 2009). The positive
in 17,18O by 3% relative to the terrestrial composition correlation between the d17,18O and d13C observed among
(Ireland et al. 2006). Hashizume and Chaussidon (2009) the organics suggests that the temperature where the
confirmed existence of both the 16O- and 17,18O-rich com- CO photodissociation occurred was higher than the
ponents among the lunar metallic grains, but concluding one for the typical molecular cloud core (10 K), which
that the 17,18O-rich component seems to be a nonsolar is the place proposed by Yurimoto and Kuramoto (2004)
component. This conclusion came from their observation where the CO self-shielding might have occurred.
that the 17,18O-rich component was more than an order of Hashizume et al. (2011) argued that the organic formation
magnitude too abundant to assign it to the solar-wind associated with the 17,18O-enrichment probably occurred
composition, whose amount was estimated from the at the envelope of the solar nebula, a slightly warm
implanted dose of solar noble gases to the lunar soil grains. (>40 K) and dense gas medium intensively illuminated
This debate may come to the end in the near-future after by the proto-Sun.
the final result for the O-isotope determination of the
solar-wind component (McKeegan et al. 2010) captured Future Directions
by the GENESIS space mission. The space-probe circled The non-mass-dependent O-isotope anomaly, broadly O
the Sun for several years to accumulate solar wind observed among the solid planetary materials, was origi-
on several substrates, which were returned to Earth nally considered to represent the origins or formation
in 2004. processes of a subset of the rocky planetary materials.
The other important end-member compositions, However, recent studies suggest that the O-isotope anom-
which may serve as tests for several models, are the pri- aly is deeply connected to the origins of the low-
mordial compositions of water ice and organic matter. temperature condensates, that is, the volatiles, represented
The models by Yurimoto and Kuramoto (2004) or Lyons by the water ice and the organic matter, which are critically
and Young (2005) predict that water ice is the carrier of the important materials in astrobiology. It is important to
17,18
O-rich isotope signature, connecting the 17,18O-rich summarize the two advantages of O-isotope over other
O atoms, first produced by the CO self-shielding in the gas light element (H, C and N) stable isotopes as a tracer.
medium, and the rocky materials at the mid-plane of the (1) Oxygen has three isotopes. We may accurately discrim-
solar-nebula, which finally trapped the isotope signature. inate the original mass-independent isotope signature
Sakamoto et al. (2007) discovered in a primitive carbona- from the mass-dependent fractionations that may occur
ceous chondrite Acfer094, a poorly characterized iron- by miscellaneous effects. (2) Oxygen is the major constit-
oxide mineral with O-isotope compositions as high as uent in both low- and high-temperature condensates. The
d17O d18O 180‰. They explained that the iron- oxygen isotope may become one of the most important
oxide mineral was produced by reactions between primor- tools to trace the entire history of the important ingredi-
dial water and the metallic Fe-Ni or iron-sulfides, and ents for a habitable planet: from their origins; their distri-
argued that their observation demonstrates the role of bution processes in the solar nebula; accretion to planets/
water ice as a carrier of the O-isotope signature produced asteroids; and the processes within the planets, namely, the
by the CO self-shielding. Hashizume et al. (2011) recently rock–water–organics interactions, possibly including the
discovered that an Antarctic carbonaceous chondrite 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
Oxygenation of the Earth’s Atmosphere O 1201
? Cyanobacteria O
Visible fossils of algae
Animals
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
1202 O Oxygenation of the Earth’s Atmosphere
evidence that O2 levels were significantly different on the environment must have acted as a powerful chemical
early Earth was the geologist Alexander MacGregor. In the buffer opposing the accumulation of atmospheric O2.
1920s, MacGregor examined Archean (pre-2.5 Ga) sedi- Consequently, we need to carefully consider the losses of
mentary rocks in modern-day Zimbabwe and deduced O2 as well as its source for understanding past changes in
that they formed in an atmosphere devoid of O2 because O2 levels. In this regard, the modern oxygen cycle helps to
they lacked the reddish-colored, oxidized iron minerals elucidate some of the basic concepts of source and sink
that are ubiquitous in analogous modern sediments. fluxes before discussing how sources and sinks were prob-
MacGregor’s conclusion greatly influenced Preston ably different on the ancient Earth.
Cloud, an American geologist, who further investigated
the geologic record for information on past atmospheric The O2 Balance Sheet: Gains and Losses
O2. Cloud also championed a suggestion originally made The theoretical subtleties of how photosynthesis can influ-
in 1959 by the marine biologist John Nursall that the ence long-term atmospheric O2 levels were not fully
relatively late appearance of macroscopic fossils animals appreciated until the 1970s, in spite of the fact that
in the geologic record reflected the slowness in reaching a French geologist, Jacques Ebelmen, correctly described
a sufficiently O2-rich atmosphere. the key ideas in 1845. Ebelmen implicitly realized that
Modern geochemical research shows that an initial photosynthesis is nearly a zero-sum activity. Oxygenic
transition to an oxygenated atmosphere occurred in the photosynthesis can be summarized by the following sche-
period 2.45–2.32 Ga, but the O2 levels attained were prob- matic equation:
ably only 0.2–2% by volume and apparently insufficient CO2 þH2 O $ CH2 OþO2 ð1Þ
for supporting macroscopic animals. This first “rise of
oxygen” is also called the Great Oxidation Event (GOE). where “CH2O” represents the average stoichiometry of
Although much less than today’s 21% by volume, the organic matter. Oxygen consumption by respiration or
post-GOE O2 levels were enough to dominate the redox decay of organic matter reverses Eq. 1, and produces no
chemistry of the atmosphere. Such O2 levels were also net oxygen on timescales greater than about 102 years.
ample for producing a stratospheric ozone (O3) layer, Oxygen only escapes respiration of oxidative decay on
which ever since has protected life on the Earth’s surface geologic timescales because a tiny fraction (0.1–0.2%) of
from biologically harmful ultraviolet radiation. The first organic carbon is buried, mainly in marine sediments,
fossils visible to the naked eye occur after the GOE at which segregates the oxygen from the organic matter.
1.87 Ga and are thought to represent a seaweed, Grypania Some organic matter is also used in biogeochemical pro-
spiralis. In a second increase in O2, over the period 750– cesses to make pyrite (FeS2) from seawater sulfate ions
580 Ma, O2 rose to levels of at least tens of percents of (SO42(aq)) as follows:
modern, that is, a concentration between 2% and present Oxygenic photosynthesis : CO2 þH2 O ¼ CH2 O þ O2 15
levels. The first animal fossils appear in the Ediacaran Period
Anaerobic decomposition : 15CH2 O þ 8SO4 2 þ 2Fe2 O3
(635–542 Ma), with the oldest soft-bodied fossils of Edia-
caran fauna at 575–565 Ma. While there is a consensus that þ 16Hþ ¼ 4FeS2 þ 15CO2 þ 23H2 O
sufficient O2 was a necessary physiological precursor to
animal life, there remains considerable debate among Net reaction : 8SO4 2 þ 2Fe2 O3 þ 16Hþ
paleontologists about whether an increase in O2 caused ¼ 15O2 þ 4FeS2 ðburiedÞ þ 8H2 O
large animals to appear or whether other evolutionary and
ð2Þ
ecological factors were crucial.
Explaining why O2 increased is more elusive than it Consequently, pyrite burial also liberates oxygen into
might first appear. We might suppose that when the atmosphere or ocean. Remarkably, Ebelmen correctly
cyanobacteria evolved, O2 would have accumulated in identified both organic carbon burial and pyrite burial as
the atmosphere. But the consensus interpretation of geo- the long-term sources of atmospheric oxygen. Unfortu-
chemical evidence (discussed below) is that cyanobacteria nately, his ideas went unnoticed and the same concepts
produced O2 for several hundred million years or had to be reinvented.
more before 2.4 Ga, while atmospheric O2 levels remained While it is useful to think of the burial of organic
<1 ppmv. A pertinent consideration is that the Earth’s carbon and pyrite as a “net source” of O2, it is important
overall chemical composition is reducing, so that reactions to appreciate that this “net” O2 is lost to numerous other
of O2 with geothermal gases in the atmosphere or certain sinks for O2 besides the reversal of eq. [1]. The geological
cations in the ocean (e.g., Fe2+) in an initially reducing sinks and sources of O2 are shown in Fig. 2. By reviewing
Oxygenation of the Earth’s Atmosphere O 1203
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)
the average composition of new sediments, American centimeter- to meter-sized animals, which require O2 to
geochemist Heinrich Holland has estimated that burial grow and breathe, have been continuously present in the
fluxes of organic carbon and pyrite produce oxygen at Phanerozoic eon (542 Ma to present) so that concentra-
rates of 10.0 3.3 Tmol O2 year1 and 7.8 4.0 Tmol tions of atmospheric O2 must have been within 0.2 0.1
O2 year1, respectively, where 1 Tmol = 1012 mol (Holland bar. The O2 source is balanced by O2 losses that include
2002). The reduction of oxidized iron from its +3 oxida- reactions in the ocean with dissolved minerals and gases
tion state to the +2 state (2Fe2O3 = 4FeO + O2) and from hot seafloor vents, and reactions of O2 dissolved in
subsequent burial also adds a minor flux of O2, whereas rainwater with continental minerals, principally those
the burial of sulfate minerals in sediments removes O2. containing elemental carbon, sulfide, and ferrous iron
Summing these burial fluxes, the total O2 net source is (Fig. 1). Gases that consume O2 in the atmosphere or
18.4 7.8 Tmol O2 year1 (Table 1). Dividing the current ocean are CH4, H2, CO, SO2, and H2S. These are released
amount of oxygen in the atmosphere-ocean reservoir either from hot rocks that melt (volcanism) or hot rocks
(3.78 1019 mol O2) by the source flux (1.8 1013 mol that do not melt (metamorphism). Estimates of their
O2 year1) gives 2.1 million years for the average fluxes are shown in Table 2. The O2 sink from submarine
amount of time an O2 molecule spends in today’s atmo- and subaerial volcanic gases is 2.3 Tmol O2 year1,
sphere-ocean system. Consequently, over such timescales, metamorphic gases consumes 3.1 Tmol O2 year1, and
the net source of O2 must be closely balanced by losses or subduction of oxidized seafloor causes a loss of 0.2 Tmol
O2 levels would wildly fluctuate. We know that 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)
15.5 6.7 O2 year1, which is estimated from the com- became more oxygenated. Let us consider evidence for the
position of average rock undergoing weathering (Table 2). GOE and Neoproterozoic increase of O2 in turn.
Thus, within the large 40% uncertainty of the data, the
source flux of O2 from organic carbon and pyrite burial of Evidence for the Great Oxidation Event
18.4 7.8 Tmol O2 year1 is matched by the total sink flux (GOE)
of 21.1 8.1 Tmol O2 year1. In the past, changes in the The different oxidation state of continental minerals that
relative proportions of source and sink fluxes evidently were formed before and after the GOE records a change
caused O2 levels to change. from an atmosphere essentially devoid of O2 to one redox
dominated by O2. The presence or absence of O2 changes
Evidence for Earth’s Oxygenation the chemistry of minerals that are formed because O2
The Great Oxidation Event at 2.45–2.32 Ga occurred in dissolves in rainwater and reacts with minerals. Three
the Paleoproterozoic era (2.5–1.6 Ga), the first of three continental indicators are red beds, paleosols, and detrital
eras within the Proterozoic eon (2.5–0.542 Ga), which grains (Holland 1984). Red beds are riverbanks, deserts,
follows the Archean eon (before 2.5 Ga). After the and floodplains with a reddish pigmentation that arises
Paleoproterozoic, the Mesoproterozoic (1.6–1.0 Ga) and when an iron oxide coating on mineral grains is produced
the Neoproterozoic (1.0 Ga–542 Ma) eras follow. The from the rusting of iron minerals. Before the GOE, no red
second rise of O2 (750–580 Ma) occurred in the beds are found in the geologic record, indicating a lack of
Neoproterozoic. We can tell that O2 levels changed by O2, whereas afterward red beds are common. Paleosols are
studying at the chemistry of rocks from these eras. The lithified soils. Before the GOE, paleosols show a loss of
Great Oxidation Event (GOE) produced binary geochem- iron, which is consistent with leaching by ancient anoxic
ical changes in the Paleoproterozoic because the atmo- rainwater, whereas after 2.2 Ga, iron is not leached. Iron
sphere changed the oxidation states (from weakly will be flushed through a soil in the form of soluble ferrous
reducing to oxidizing), whereas the Neoproterozoic iron ions (Fe2+) if rainwater has little dissolved O2 but if
changes are subtler because an oxic atmosphere merely the air contains >0.2% O2, iron will be oxidized into
Oxygenation of the Earth’s Atmosphere O 1205
insoluble and immobile oxides such as ▶ hematite was (microbially) oxidized and precipitated as iron min-
(Fe2O3). Cerium is another redox-sensitive element that erals in the BIFs. In the Paleoproterozoic, BIFs decline in
has also been examined in paleosols. Cerium oxidizes abundance and disappear after 1.8 Ga. Chromium (Cr)
from Ce3+ to Ce4+ to form cerianite (CeO2), so the pres- isotopes in the BIFs also record changes in atmospheric
ence of Ce3+-rich minerals in paleosols from the Archean O2. When insoluble Cr3+ is oxidized to soluble Cr6+ dur-
implies an early anoxic atmosphere. Detrital grains in ing oxidative weathering, the heavy isotope 53Cr is prefer-
sediments are those that are never completely dissolved entially oxidized relative to 52Cr. After transport from
during the process of weathering. Rounded detrital grains rivers to the ocean, the Cr isotopic fractionation is pre-
from sediments deposited by turbulent Archean rivers served in ocean sediments. An increase in the abundance
commonly contain minerals that would only survive at of 53Cr after 2.4 Ga compared to Archean samples further
very low levels of O2. Detrital grains of pyrite (FeS2), confirms the GOE.
uraninite (UO2), and siderite (FeCO3) place bounds on The strongest evidence for the GOE is a large amount
the concentration of Archean O2 of roughly <2%, <0.2%, of mass-independent fractionation (MIF) of ▶ sulfur iso-
and <0.02%, respectively, compared to today’s 21% O2. topes in sedimentary rocks older than 2.45 Ga compared
In modern oxygenated waters, uraninite dissolves to to little or no sulfur isotope MIF in rocks younger than
form soluble U6+ ions, pyrite oxidizes to soluble sulfate 2.32 Ga. In modern rocks, 32S, 33S, and 34S obey “mass-
(SO42) and hematite, and siderite rusts to produce insol- dependent” fractionation, in which the difference in abun-
uble hematite. dance between 33S and 32S is approximately half that
The chemistry of seawater also changed after the GOE, between 34S and 32S, that is, a linear relationship with
with increased levels of sulfate and much decreased levels mass difference. Most processes, such as microbial sulfate
of redox-sensitive elements such as manganese and iron reduction, produce mass-dependent isotope fraction-
that are insoluble when oxidized. Marine sulfur isotopes ation. In contrast, mass-independent fractionation in sul-
are the data that indicate an increase in sulfate concentra- fur isotopes is where the relative abundance of different
tions. Today, sulfate-reducing bacteria produce most of isotopes deviates from linear, mass-dependent fraction-
the sulfide in marine sediments. These bacteria reduce ation. Laboratory studies show that large MIF is produced
32
SO42 in preference to 34SO42 so that bacterial sulfide exclusively from photochemical reactions, such as the
is enriched in 32S. However, this fractionation ceases in photolysis of SO2. Three conditions are needed for the
waters with <0.2 mM sulfate concentration. Archean sul- production and preservation of the sulfur isotope MIF: O2
fides have 34S/32S ratios close to an unfractionated mantle levels below about 1 ppmv, sufficient sulfur gas in the O
value, implying Archean oceans with <0.2 mM sulfate, atmosphere, and relatively large concentrations of reduc-
compared to 28.9 mM in today’s surface seawater. Rivers ing gases. In a high-O2 atm, sulfur gases are rapidly oxi-
ultimately supply sulfate to the ocean by washing sulfate dized and rained out as dissolved sulfate. But in the
off the land where it is produced by the oxidation of absence of O2 and in the presence of reducing gases such
sulfides by oxygenated rainwater. Consequently, the low as 102–103 ppmv methane, sulfur exits the atmosphere as
concentration of seawater sulfate before the GOE is con- sulfide, elemental sulfur, or sulfate, which allows the mass-
sistent with low atmospheric O2. By 2.3–2.2 Ga, sulfides independent fractionation produced by anoxic photo-
with significant 34S depletions formed ubiquitously in the chemistry to be segregated and preserved in different
ocean, reflecting increased sulfate and the rise of O2. When sedimentary minerals. In summary, the sulfur isotope
O2 rose, it also oxidized and titrated manganese out of the MIF provides evidence for an Archean atmosphere with
ocean, producing the world’s largest manganese deposit both low O2 and high methane.
found in the 2.2 Ga Hotazel Formation in South Africa. Donald Canfield (of the University of Southern
The presence of ▶ banded iron formations (BIFs) pro- Denmark) has proposed that a significant fraction of the
vides evidence that the deep ocean was devoid of O2 in deep ocean remained sulfidic rather than oxygenated dur-
the Archean, unlike today. By definition, BIFs are lami- ing much of the Proterozoic, after 1.8 Ga (Canfield 2005;
nated marine sedimentary rocks containing 15 wt% Lyons et al. 2009). In this model, BIFs declined because
iron, with alternating iron-rich and iron-poor layers. The increased sulfate in the surface ocean caused sulfate-
iron mainly originated from hydrothermal sources in the reducing bacteria to provide a flux of sulfide to the deep
deep ocean, such as midocean ridges. Today, iron is oxi- ocean, which scavenged ferrous iron into insoluble iron
dized in the deep ocean and deposited on the flanks of the sulfides. This is an alternative to a model where the disap-
ridges. But in the anoxic depths of Archean oceans, ferrous pearance of BIFs at 1.8 Ga is explained by the oxygenation
iron was transported to the continental shelves where it 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 stromatolites, redox-sensitive metals, and sulfur isotopes,
seafloor is euxinic, meaning oxygen free with H2S. Molyb- which we discuss in turn.
denum (Mo) isotopes are useful indicators of oxygenation Biomarkers are recognizable derivatives of biological
and euxinia because oxic uptake of Mo on marine solids molecules, which are usually in the form of certain hydro-
preferentially removes 95Mo relative to 98Mo, leaving the carbons found in kerogen (noncrystalline sedimentary
ocean isotopically heavy. In contrast, in times of low organic matter) or oil. Chemical changes induced when
oxygenation, marine Mo is isotopically light. In highly sediments convert into rock strip biological molecules of
euxinic deep waters, Mo is scavenged efficiently and functional groups but can often leave a distinctive molec-
tends to capture the Mo isotope composition of seawater. ular backbone, in a way that is analogous to the more
Rocks that are 95Mo-enriched from the 1.7–1.4 Ga familiar macroscopic process that removes flesh to leave
MacArthur Basin in Australia suggest that the deep ocean fossilized skeletons that we can associate with particular
in the mid-Proterozoic seafloor was commonly oxygen- life forms. Biomarkers from the Archean have been the
poor, with a greater proportion of euxinia than today. The subject of controversy because of the possibility of con-
data suggest that the euxinia was probably only several tamination from modern molecules or fluid migration
percent of the ocean’s area, but this is enough to have had over geologic time and uncertainties regarding their
a significant effect in depleting trace metals in the specificity for particular organisms and metabolisms.
Mesoproterozoic Ocean, which may have impacted marine Recent advances in ultraclean drilling and the ability to
ecology for a billion years until the Neoproterozoic. analyze oil-bearing fluid inclusions help to address con-
cerns about contamination. Biomarkers for cyanobacteria
Evidence for a Neoproterozoic “Second Rise” are 2-alpha-methylhopanes, which are derived from com-
in Oxygen pounds in cell membranes. Long chain varieties of these
Sulfur and chromium isotopes in marine sediments indi- particular molecules (i.e., >C31) occur very rarely outside
cate that O2 increased in the late Neoproterozoic. At the cyanobacteria. In addition, steranes derive from ste-
this time, marine sedimentary sulfides occur with rols, which are found in the membranes of eukaryotes and
32
S-enrichment exceeding the isotope discrimination synthesized in a pathway that requires free molecular
of sulfate-reducing bacteria. The interpretation is that oxygen. For example, 11 molecules of O2 are required to
isotopically light sulfide was re-oxidized at the sediment- produce one molecule of cholesterol. No bacteria are
water interface and cyclically re-reduced by sulfur- known to synthesize C26–C30 sterols and archaea do not
disproportionating bacteria, which increased the isotope synthesize sterols at all. The simultaneous co-detection of
fractionation. The greater sulfur isotope fractionation is long chain (>C31) 2-alpha-methylhopanes with eukary-
therefore related to oxygenation of the near-shore seafloor. otic sterols is thus a reasonable biosignature for oxygenic
Canfield (2005) has estimated that the oxygenation photosynthesis. Such a combination of biomarkers has
required atmospheric O2 concentrations in excess of been found in fluid inclusions in the 2.45 Ga Matinenda
1–4%. A large increase in the abundance of marine 53Cr Formation at Elliot Lake, Canada, which consists of sand-
relative to 52Cr after 750 Ma also confirms an increase in stones deposited prior to the GOE (Dutkiewicz et al.
atmospheric O2. Oxygen would have mobilized greater 2006). Also, meticulous measurements of biomarkers
amounts of chromium during continental oxidative from 2.72 to 2.56 Ga rocks in the Hamersley Province
weathering, which is the mechanism of isotopic fraction- of northwest Australia (Eigenbrode et al. 2008) and
ation mentioned earlier. Fossil evidence for animals 2.67–2.46 Ga rocks from the Transvaal of South Africa
greater than a few millimeters or more in size also suggests (Waldbauer et al. 2009) suggest the presence of microbial
that O2 concentrations exceeded 2%, which is the mini- ecosystems that produced O2 as well as consumed it.
mum level that can support aerobic metabolism in such Stromatolites (laminated rocks formed by microbes)
animals via O2 supplied by diffusion. from a 2.72 Ga lake in the Tumbiana Formation in north-
west Australia possibly confirm the presence of O2
When Did Oxygenic Photosynthesis Evolve? production. Geologic evidence shows that the lake
In order to understand the early history of atmospheric lacked sulfate and hydrothermal reductants needed
O2, we need to examine the evidence for the advent of for non-oxygen-producing photosynthesis. Consequently,
oxygenic photosynthesis. In particular, the geologic record a process of elimination suggests that microbial
suggests that oxygenic photosynthesis arose 300 Ma or communities using oxygenic photosynthesis may
more before the GOE. This timing is inferred from have constructed these particular stromatolites (Sakurai
a variety of proxies including biomarkers, nonmarine et al. 2005).
Oxygenation of the Earth’s Atmosphere O 1207
Furthermore, the mobility of redox-sensitive metals flux of volatile and aqueous reductants, that is, chemicals
indicates the presence of some O2 before the GOE, and that consume oxygen, scavenged the O2. In time, the
therefore the presence of oxygenic photosynthesis. Molyb- geological sources of reductants declined to the point
denum (Mo), rhenium (Re), and chromium (Cr) are where the O2 flux produced from organic carbon burial
soluble only when oxidized. Consequently, a spike in Mo exceeded the kinetically rapid consumption flux. At this
and Re concentrations found in 2.5 Ga sedimentary rocks tipping point, oxygen flooded the atmosphere until O2
in the Hamersley district of northwest Australia suggests levels reached a new plateau where continental weathering
that a transient “whiff ” of O2 occurred in the atmosphere became the predominant balance for O2 production.
about 100 million years prior to the GOE (Anbar et al. Various suggestions have been made for why the propor-
2007). The whiff is confirmed by contemporaneous evi- tion of reductants in volcanic or metamorphic outgassing
dence of sulfur isotopic fractionation indicating a spike of might diminish and cause such a tipping point. One
sulfate. Enrichments of 53Cr relative to 52Cr (which are suggestion is that a change in the style of outgassing
induced by oxidative weathering) are also found in banded from submarine to subaerial volcanism favored a decline
iron formations at low levels for at least 300 million years in the proportion of reductants. Another consideration is
prior to the GOE, which is consistent with the biological that an anoxic atmosphere would always allow a relatively
production of O2 (Frei et al. 2009). In summary, multiple high concentration of hydrogen-bearing reducing gases.
lines of evidence suggest that oxygenic photosynthesis At high altitude, the decomposition of such gases causes
existed at least 0.3 billion years before the GOE. hydrogen to escape to space at a rate that is significant on
geologic timescales, which oxidizes the whole Earth and
Theories for Oxygenation diminishes further release of reductants. In particular, in
At present, a framework has been developed for under- an anoxic atmosphere, methane reaches concentrations
standing why the GOE occurred, but the cause of the hundreds of times greater than today’s 1.8 ppmv. Most
second rise of O2 in the Neoproterozoic remains very methane is ultimately produced microbially from fermen-
obscure (Catling and Claire 2005). tation of photosynthesized organic matter. Ultraviolet
The GOE happened because either O2 production light decomposes methane in the upper atmosphere, caus-
increased or O2 consumption declined. The ancient, pre- ing it to lose hydrogen. When hydrogen escapes from the
2.45 Ga oceans were largely devoid of sulfate, so that planet, oxygen is gained irreversibly through a schematic
organic carbon burial dominated O2 production prior to reaction as follows:
the GOE, while pyrite burial was unimportant. Conse- O
CO2 þ H2 O þ biology ! CH4 þ O2
quently, it has been argued that O2 rose because the rate
of organic carbon burial (O2 production) increased, either þ photochemistry ! 4Hðlost to spaceÞ þ CO2 þ O2
as a pulse or long-term trend. One suggestion is that the ð3Þ
growth of early continental shelves promoted organic
carbon burial and oxygen production. But there are Evidence for this hypothesis is found in the redox
grounds for skepticism about this proposal. Organic mat- components of the crust. If photosynthesis were the only
ter extracts the light carbon isotope, 12C, from seawater, process responsible for segregating redox components, the
but marine carbon, recorded in ancient limestone, number of moles of equivalent O2 in the Earth’s crust
does not significantly decrease in 12C content between should be balanced by equimolar reduced carbon
mid-Archean and Mesoproterozoic rocks. Notably, at (Eq. 1). But instead there is excess oxygen in the crust by
2.3–2.1 Ga, many marine carbonates are unusually a factor of 1.5–2.2 (Catling and Claire 2005; Sleep 2005).
depleted in 12C relative to 13C, which could be explained The excess can be quantitatively reconciled with time-
if there was a large pulse of burial of organic carbon, but integrated hydrogen escape and net oxidation expected
then the 12C content returns to its previous value. A burial from a methane-rich Archean atmosphere.
pulse would merely generate a parallel pulse of O2, given Various geological and biological causes have been
the short residence time of O2 and not a persistently suggested for the Neoproterozoic second rise of oxygen,
increased level of O2. Instead, the pulse was more plausibly but the problem remains unsolved. Geological proposals
an effect of increased O2 because the Paleoproterozoic include the enhanced production of clays that adsorbed
excursions in the 12C of limestones follow the rise of O2 and buried organic matter, or the assembly of
before 2.32 Ga. a supercontinent whose weathering added nutrients to
A widely accepted explanation for why little atmo- the sea and stimulated organic carbon burial. Also, it has
spheric O2 accumulated in the Archean is that a large been suggested that moderately high levels of methane
1208 O Oxygenation of the Earth’s Atmosphere
Definition
Oxygenic Phototrophic Bacteria The ▶ ozone layer refers to the ozone within the strato-
sphere of the Earth, where over 90% of the Earth’s ozone
▶ Cyanobacteria, Diversity and Evolution of 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
Oxyphotobacteria The existence of an ozone layer in the upper regions of our
atmosphere was detected in 1913 by Charles Fabry and
▶ Cyanobacteria, Diversity and Evolution of 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
Ozone measure of the columnar density of ozone above the
Earth surface.
Definition
Ozone is an important trace constituent of the stratosphere Overview
where it forms the ozone layer. It consists of three atoms of On Earth, the stratospheric ozone layer is found at an
oxygen arranged as an open triangle. Ozone is produced altitude from 20 to 40 km. It is created when ultraviolet
in the three-body reaction O þ O2 þ M = O2 þ M, radiation from the Sun strikes the oxygen molecules in the
where M represents a stabilizing molecule (N2 or O2) stratosphere. By these photochemical processes oxygen
and O is produced by the photochemical dissociation molecules (O2) are dissociated to atomic oxygen (O).
of O2. Some other components (NO, Cl, CFCs) catalyze The atomic oxygen quickly combines with further oxygen
the reaction of ozone destruction O3 þ O = 2O2. molecules to form ozone (O3). Ozone can be split again
1210 O Ozone Layer