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Principles of terrestrial ecosystem ecology.pdf

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232 10. Aquatic Carbon and Nutrient Cycling<br />

Frequency<br />

15<br />

10<br />

5<br />

0<br />

15<br />

10<br />

5<br />

0<br />

Marine phytoplankton<br />

Freshwater phytoplankton<br />

10 100<br />

N:P (by atoms) [log scale]<br />

caused phytoplankton blooms large enough to<br />

be seen from satellites, indicating that iron<br />

limits the capacity <strong>of</strong> phytoplankton to use<br />

nitrogen and phosphorus. During glacial<br />

periods there may have been 10-fold greater<br />

input <strong>of</strong> iron- and phosphorus-bearing dust to<br />

the oceans, thus stimulating ocean productivity<br />

and in turn lowering atmospheric CO2 concentrations<br />

(Falkowski et al. 1999). The key role <strong>of</strong><br />

iron in regulating production in some sectors <strong>of</strong><br />

the open ocean has led to the suggestion that<br />

large-scale iron fertilization might stimulate<br />

ocean production sufficiently to scavange large<br />

amounts <strong>of</strong> CO2 from the atmosphere and<br />

sequester it in deep oceans in the form <strong>of</strong> dead<br />

organic matter. The iron-addition experiments,<br />

however, show that this stimulation <strong>of</strong> production<br />

is relatively short-lived, presumably<br />

because other elements quickly become limiting<br />

to production, as soon as the iron demands<br />

<strong>of</strong> phytoplankton are met. Grazing is another<br />

factor that contributes to low phytoplankton<br />

biomass and productivity in portions <strong>of</strong> the<br />

open ocean. In some HNLC zones <strong>of</strong> the ocean<br />

there is simply not enough phytoplankton<br />

biomass to use the nutrients that are available<br />

(Valiela 1995).<br />

N limited<br />

P limited<br />

No data available<br />

1 1000<br />

Figure 10.7. Frequency distribution<br />

<strong>of</strong> the N : P ratio in marine and<br />

fresh-water phytoplankton. Nutrient<br />

addition experiments (shaded<br />

bars) indicate that the high N:P<br />

ratios in lakes reflect phosphorus<br />

limitation <strong>of</strong> algal growth, whereas<br />

the generally low N:P ratios <strong>of</strong><br />

marine phytoplankton commonly<br />

reflect nitrogen limitation. (Modified<br />

with permission from Springer-<br />

Verlag; Valiela 1995.)<br />

The strong nutrient limitation and lack <strong>of</strong><br />

CO2 limitation <strong>of</strong> productivity in most <strong>of</strong> the<br />

world’s oceans make it unlikely that marine<br />

productivity will respond directly to increasing<br />

atmospheric CO2. Nutrient limitation <strong>of</strong> marine<br />

production, however, makes these <strong>ecosystem</strong>s<br />

potentially vulnerable to anthropogenic<br />

nutrient inputs. Estuaries, for example, have<br />

been substantially altered by the large nutrient<br />

inputs from agricultural lands. Run<strong>of</strong>f and<br />

sewage effluents have substantially altered<br />

coastal <strong>ecosystem</strong>s near heavily populated<br />

or agricultural areas (Howarth et al. 1996).<br />

Oxygen depletion by decomposers in the water<br />

column <strong>of</strong> the Mississippi Delta, for example,<br />

has created a large dead zone in the Gulf <strong>of</strong><br />

Mexico. Although the impacts <strong>of</strong> nutrients on<br />

the open ocean may be more subtle and difficult<br />

to detect because most pollution sources<br />

are remote, they could, over the long term, be<br />

important because <strong>of</strong> the high degree <strong>of</strong> nutrient<br />

limitation <strong>of</strong> pelagic <strong>ecosystem</strong>s and their<br />

large aerial extent.<br />

Ocean productivity is ultimately limited by<br />

the rate <strong>of</strong> nutrient supply from the land or<br />

deep ocean waters. For this reason, productivity<br />

is greater in coastal waters than in the

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