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Principles of terrestrial ecosystem ecology.pdf

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Nitrogen output (g m -2 yr -1 )<br />

5<br />

4<br />

3<br />

2<br />

1<br />

0<br />

North America<br />

Europe<br />

0 1 2 3 4<br />

Nitrogen deposition (g m -2 yr -1 )<br />

Figure 9.9. Comparisons <strong>of</strong> inputs from nitrogen<br />

deposition and nitrogen outputs in solution from<br />

forests <strong>of</strong> North America and Europe. There is a<br />

strong relationship between inputs and outputs in<br />

nitrogen-saturated <strong>ecosystem</strong>s. (Data from Tietema<br />

and Beier 1995, Fenn et al. 1998.)<br />

<strong>of</strong>ten has nitrate concentrations that exceed<br />

public health standards.<br />

Nitrogen leached from <strong>terrestrial</strong> <strong>ecosystem</strong>s<br />

moves in groundwater to lakes and rivers. The<br />

movement <strong>of</strong> nitrate to the North Atlantic<br />

Ocean from major rivers has increased 6- to 20fold<br />

in the past century (Howarth et al. 1996),<br />

primarily due to increased inputs <strong>of</strong> fertilizer,<br />

atmospheric deposition, nitrogen fixation by<br />

crops, and food imports (see Chapter 15).<br />

Nitrate in coastal marine systems frequently<br />

increases productivity and detritus accumulation.<br />

The resulting stimulation <strong>of</strong> decomposition<br />

can reduce oxygen concentrations<br />

sufficiently to kill fish, particularly in winter,<br />

when primary production is temperature<br />

limited (see Chapter 14). The nitrogen loading<br />

from agricultural and urban systems in the<br />

Mississippi drainage, for example, has produced<br />

a “dead zone” where this river enters the<br />

Gulf <strong>of</strong> Mexico (Mitsch et al. 2001).<br />

Solutions that move through the soil must<br />

maintain a balanced charge, with negatively<br />

charged ions like nitrate balanced by cations.<br />

Therefore, every nitrate ion that leaches from<br />

soil carries with it a cation such as calcium,<br />

5<br />

Other Element Cycles 215<br />

potassium, and ammonium to maintain charge<br />

balance. When cation loss by leaching exceeds<br />

the rate <strong>of</strong> cation supply by weathering plus<br />

deposition, the net loss <strong>of</strong> cations can lead to<br />

cation deficiency (Driscoll et al. 2001). After<br />

these nutrient cations are depleted, nitrate<br />

takes with it H + and/or Al 3+ , which are deleterious<br />

to downstream <strong>ecosystem</strong>s. Nitrification<br />

also generates acidity:<br />

2NH4 + + 3O2 Æ 2NO2 - + H2O + H + (9.5)<br />

The hydrogen ion released in this reaction<br />

exchanges with other ions on cation exchange<br />

sites in the soil, making these cations more vulnerable<br />

to leaching loss.<br />

Erosional Losses<br />

Erosion is a natural pathway <strong>of</strong> nitrogen loss<br />

that <strong>of</strong>ten increases dramatically after land use<br />

changes. As with leaching, erosional losses <strong>of</strong><br />

nitrogen include both organic and inorganic<br />

forms, although organic forms associated with<br />

soil aggregates and particles are most important.<br />

In some <strong>ecosystem</strong>s, especially those on<br />

unstable slopes or in areas exposed to high<br />

winds, erosion is a dominant natural pathway <strong>of</strong><br />

nitrogen loss.<br />

Other Element Cycles<br />

Phosphorus<br />

Weathering <strong>of</strong> primary minerals is the major<br />

source <strong>of</strong> new phosphorus to <strong>ecosystem</strong>s. In<br />

contrast to nitrogen, whose major source is the<br />

atmosphere, phosphorus enters <strong>ecosystem</strong>s primarily<br />

by weathering <strong>of</strong> rocks (Fig. 9.10). The<br />

weathering <strong>of</strong> phosphorus-containing apatite<br />

by the carbonic acid generated from soil respiration,<br />

for example, releases phosphorus in<br />

available forms (Eq. 9.6) that can be taken up<br />

directly by plants or microorganisms or be<br />

adsorbed or precipitated (see Chapter 3).<br />

Ca5(PO4)3 + 4H2CO3 Æ<br />

5Ca 2+ + 3HPO4 2- + 4HCO3 + H2O (9.6)<br />

Phosphorus inputs from weathering depend<br />

on the mineralogy <strong>of</strong> the parent material, the

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