go extinct survived by retreating, for example, to
anaerobic environments, such as deep in crustal
rocks, organic-rich muds, and animal digestive tracts,
where they continue to thrive today. New forms of
life emerged with superoxide dysmutase and catalase
enzymes, which catalyze the reduction of oxic free
radicals to water. Finally, another group (the
Eukarya; Figure 7) developed aerobic metabolism,
providing energy yields approximately twenty times
greater than those of anaerobic metabolism (see Pre-
cambrian: Eukaryote Fossils). The mitochondria –
cell organelles contained in most Eukarya, which fa-
cilitate oxidative metabolism – arose from the endo-
symbiosis of proteobacteria. These organelles
actually contain their own subunits of DNA that
further implicate a eubacterial heritage.
The increased energy yield of aerobic metabolism
set the stage for the evolution of all life above the
unicellular level. The energy source that supports
the aerobic biosphere is the Sun – thus established
oxygenic photosynthesis and aerobic respiration
governed the flow of carbon and oxygen through
the atmosphere–hydrosphere system. During the Ar-
chaean the atmosphere and hydrosphere initially con-
tained small amounts of free oxygen from photolysis
of water vapour and then increasing amounts from
oxygenic photosynthesis. By the end of the Archaean–
Early Proterozoic, oxygen levels had begun to creep
upwards, balanced by oxygen-consuming reactions
such as weathering, hydrothermal activity, respir-
ation, oxidation of organic matter, and differential
rates of organic-matter burial. By about 2 Ga, levels
of oxygen were about 1% of PAL. A transition era
ensued, with oxygen levels fluctuating around the 1%
PAL value and the oceans remaining reducing and
sulphidic. After about 1.8 Ga, oxygen-consuming re-
actions were generally exhausted and free-oxygen
concentrations reached about 10% of PAL. Over
time, these levels stabilized at near ‘normal’ Phaner-
ozoic atmospheric concentrations. It remains unquan-
tified how rapidly oxygen levels increased during the
Proterozoic.
The rise in atmospheric oxygen had an acute effect
on the surface environment, not only because of its
toxicity to many microbial organisms (providing the
motive force to drive the evolution of more efficient
aerobic metabolisms) but also by establishing an ef-
fective ultraviolet screen. Ozone is far more effective
than diatomic oxygen at absorbing ultraviolet. The
ozone screen formed in the stratosphere from accu-
mulating O
2
that photodissociated to produce free
oxygen radicals (O
), which then recombined with
O
2
to make ozone. This ozone screen effectively
made dry land habitable for plants and animals
by the Palaeozoic. Industrial pollutants such as
chlorofluorocarbons are now severely damaging the
ozone layer.
A Neoproterozoic Snowball Earth
The increased oxidation of the surface zone in the
Middle Proterozoic was probably a consequence of
the sequestration of large quantities of organic carbon
in sediments. From the oxygenic photosynthesis reac-
tion, it can be seen how this scenario leads to a net loss
of carbon dioxide and a net increase of oxygen in the
atmospheric reservoir. The boosted greenhouse effect
of heightened levels of carbon dioxide before about
2.5 Ga was lost to the Earthat a most inopportune time.
Levels of solar luminosity were about 10%–15%
lower at 2.2 Ga than at present (Figure 1). Loss of
insolation from the carbon dioxide greenhouse
spelled disaster for the Proterozoic Earth. The planet
froze over, locking the oceans in ice and creating a
high-albedo feedback loop that kept the planet frozen
for extended periods, until levels of carbon dioxide
increased due to passive volcanism and outgassing.
Heightened levels of carbon dioxide warmed the at-
mosphere and dark dusty ice reduced the albedo,
causing catastrophic melting and massive planetary
warming. Subsequent weathering in a carbon diox-
ide-rich atmosphere, combined with massive algal
blooms, led to enhanced burial of organic carbon,
drawdown of carbon dioxide levels, and a renewed
snowball Earth. The cycle is thought to have been
broken by the secular increase in solar luminosity
and a steady redistribution of the continents via
plate motions. The survivors of these repeated ‘ice-
house’ and ‘hothouse’ Earths were multicellular or-
ganisms that inherited a more stable environment
high in oxygen (see Palaeoclimates).
The Phanerozoic Atmosphere
Phanerozoic Atmospheric Changes
Geochemical evidence from the study of palaeosols,
coupled with data from carbonate and organic
carbon in sediments as well as sedimentary pyrite
and the chemistry of sedimentary silicate minerals,
has been used to improve models of the carbon cycle
of palaeoenvironments. These models have been used
to document and explain fluctuations in levels of
oxygen and carbon dioxide over Phanerozoic time.
Long-term changes in the carbon dioxide and oxygen
concentrations in the Phanerozoic atmosphere are
summarized in Figure 8.
In the first part of the Phanerozoic, the Early
Palaeozoic (Cambrian–Ordovician), evidence indi-
cates that levels of carbon dioxide were about fifteen
times PAL. These declined to within a few percent of
204 ATMOSPHERE EVOLUTION