Many organisms catalyze the precipitation of
CaCO
3
from seawater through active biomineraliza-
tion and shell formation; for example, foraminifera,
coccoliths, corals, and shellfish contribute to the per-
manent removal of atmospheric CO
2
. Marine organ-
isms generally form their shells at relatively shallow
depths and sink to the shelf floor or abyssal depths after
death. In regions where the water depth is no greater
than about 1 km, the shells reach the seafloor more or
less intact because these waters are saturated with re-
spect to calcite and aragonite. However, at greater
depths, where pressures are greater, temperatures are
lower, and CO
2
concentrations are elevated, seawater
is far more corrosive to CaCO
3
. The level at which the
rate of carbonate dissolution balances the downward
flux of carbonate settling to the seafloor is called the
carbonate compensation depth (CCD). This depth
varies from ocean to ocean and has varied over time
in response to carbonate productivity, temperature,
atmospheric CO
2
, and ocean circulation changes.
Geological Evolution of the Global
Carbon Cycle
Many scientists suspect that the greenhouse effect
and, in particular, the level of CO
2
in the atmosphere
were different in the geological past. One major
reason for this suspicion derives from the ‘Faint
Young Sun Paradox’. Because the sun is probably
heating up over time – owing to the increasingly
exothermic nuclear fusion of lighter elements, form-
ing heavier elements in the sun’s central core – it is
probable that the input of solar energy to Earth was
significantly lower in the past, by as much as 1% for
every 100 million years, according to some estimates.
However, Earth’s sedimentary record provides evi-
dence for the existence of liquid water at least 4 bil-
lion years ago, indicating that surface temperatures
were not much lower then than they are now. There-
fore, it appears that, over time, Earth’s atmosphere
has become increasingly less efficient in retaining
solar energy. Model calculations indicate that, were
CO
2
the only relevant greenhouse gas, CO
2
levels
must have been as much as 10 000 times higher
during the Early Precambrian. However, because me-
thane is likely to have been a major greenhouse gas
during early Earth history, before the increase of at-
mospheric oxygen levels, such estimates are likely to
represent maximum CO
2
concentrations only.
p0060 On geological time-scales, atmospheric CO
2
levels
are thought to be regulated by negative feedbacks
between climate and silicate weathering rates.
First, increased temperatures due to higher CO
2
levels
would accelerate chemical weathering rates, thus
effectively slowing any CO
2
increase. Second,
weathering requires rainfall, which would most likely
increase as temperatures rise, due to the acceleration
of the hydrological cycle. Third, enhanced CO
2
fertil-
ization of plant growth would also help to increase
weathering rates by stabilizing soils and encour-
aging chemical weathering over physical weathering.
This does not mean, however, that CO
2
levels have
remained constant over geological time. Chemical
weathering rates are imperfectly tied to CO
2
levels,
being related also to additional parameters, includ-
ing tectonics, vegetation cover, and palaeogeography,
whereas atmospheric CO
2
may be influenced by
independent changes in the input and output of
CO
2
. In the geological past, higher CO
2
levels could
be sustained because chemical weathering rates on
the continents were lower in the absence of deep
soils, before the introduction of vascular plants in
the Devonian period. A higher PCO
2
was also made
possible by higher volcanic outgassing rates of mantle
CO
2
and the abiotic nature of much carbonate depos-
ition. Before the introduction of pelagic carbonate
producers, such as planktic foraminifera by the Juras-
sic, carbonate deposition would have been restricted
to shallow shelf environments, thus increasing calcite
saturation levels and rendering deposition rates
vulnerable to changing sea-level (palaeogeography).
Although the introduction of land plants in the
Devonian period does not appear to have had an
immediate effect on climate (by sequestering more
atmospheric CO
2
), the deposition of massive peat
deposits in shallow coastal environments during the
Carboniferous and Permian periods almost certainly
did. Long-term (10
7
years) increases in organic carbon
burial during this interval are thought to have re-
duced CO
2
levels by as much as 70% (Figure 6),
causing a series of glaciations, while allowing oxygen
to build up in the atmosphere. One consequence
of higher O
2
is to increase air pressure; it seems
plausible that the well-recorded insect gigantism
during this interval was related to the added buoy-
ancy provided by greater air pressures. Such long-
term, but ultimately unsustainable, imbalances in
the proportion of carbon buried as organic carbon
relative to carbonate carbon may have caused changes
to both the CO
2
and the O
2
atmospheric budgets at
other times, too, but the carbon isotopic record sug-
gests that this proportion has been more or less con-
stant at 1:4, respectively, on >10
7
year time-scales
since 3 Ga. Geochemical models show that CO
2
levels
during the Phanerozoic were also higher than at
present (Figure 6), and reached peaks during the
Early Phanerozoic and more recently during the Cret-
aceous period when high PCO
2
is likely to have con-
tributed to the equable and balmy ‘Greenhouse’
climates typical of that interval in Earth history.
Despite the existence of regulatory feedbacks on
atmospheric CO
2
, the Earth system is still susceptible
340 CARBON CYCLE