Principles of terrestrial ecosystem ecology.pdf

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Nitrogen output (g m -2 yr -1 ) 5 4 3 2 1 0 North America Europe 0 1 2 3 4 Nitrogen deposition (g m -2 yr -1 ) Figure 9.9. Comparisons of inputs from nitrogen deposition and nitrogen outputs in solution from forests of North America and Europe. There is a strong relationship between inputs and outputs in nitrogen-saturated ecosystems. (Data from Tietema and Beier 1995, Fenn et al. 1998.) often has nitrate concentrations that exceed public health standards. Nitrogen leached from terrestrial ecosystems moves in groundwater to lakes and rivers. The movement of nitrate to the North Atlantic Ocean from major rivers has increased 6- to 20fold in the past century (Howarth et al. 1996), primarily due to increased inputs of fertilizer, atmospheric deposition, nitrogen fixation by crops, and food imports (see Chapter 15). Nitrate in coastal marine systems frequently increases productivity and detritus accumulation. The resulting stimulation of decomposition can reduce oxygen concentrations sufficiently to kill fish, particularly in winter, when primary production is temperature limited (see Chapter 14). The nitrogen loading from agricultural and urban systems in the Mississippi drainage, for example, has produced a “dead zone” where this river enters the Gulf of Mexico (Mitsch et al. 2001). Solutions that move through the soil must maintain a balanced charge, with negatively charged ions like nitrate balanced by cations. Therefore, every nitrate ion that leaches from soil carries with it a cation such as calcium, 5 Other Element Cycles 215 potassium, and ammonium to maintain charge balance. When cation loss by leaching exceeds the rate of cation supply by weathering plus deposition, the net loss of cations can lead to cation deficiency (Driscoll et al. 2001). After these nutrient cations are depleted, nitrate takes with it H + and/or Al 3+ , which are deleterious to downstream ecosystems. Nitrification also generates acidity: 2NH4 + + 3O2 Æ 2NO2 - + H2O + H + (9.5) The hydrogen ion released in this reaction exchanges with other ions on cation exchange sites in the soil, making these cations more vulnerable to leaching loss. Erosional Losses Erosion is a natural pathway of nitrogen loss that often increases dramatically after land use changes. As with leaching, erosional losses of nitrogen include both organic and inorganic forms, although organic forms associated with soil aggregates and particles are most important. In some ecosystems, especially those on unstable slopes or in areas exposed to high winds, erosion is a dominant natural pathway of nitrogen loss. Other Element Cycles Phosphorus Weathering of primary minerals is the major source of new phosphorus to ecosystems. In contrast to nitrogen, whose major source is the atmosphere, phosphorus enters ecosystems primarily by weathering of rocks (Fig. 9.10). The weathering of phosphorus-containing apatite by the carbonic acid generated from soil respiration, for example, releases phosphorus in available forms (Eq. 9.6) that can be taken up directly by plants or microorganisms or be adsorbed or precipitated (see Chapter 3). Ca5(PO4)3 + 4H2CO3 Æ 5Ca 2+ + 3HPO4 2- + 4HCO3 + H2O (9.6) Phosphorus inputs from weathering depend on the mineralogy of the parent material, the
216 9. Terrestrial Nutrient Cycling N 2 E E E N org NH 4 + S 2- org SO4 Ca org Rock Rock Ca 2+ NO 3 - NH 3 2- SO4 H 2 S N2 N2O NO x Figure 9.10. Comparison of natural element cycles with respect to the relative importance of internal recycling, inputs, and outputs. Inputs of nitrogen come primarily from the atmosphere, whereas inputs of phosphorus, potassium, calcium, and most unessential elements such as lead (Pb) come primarily from rocks. Sulfur comes from both the E E E Rock Rock P org Rock K + Pb PO 4 3- P bound atmosphere and rocks. Over long time scales, atmospheric inputs of all elements can be important. Element losses occur through leaching, erosion (E), and, in the case of nitrogen and sulfur, gaseous emission. Subscripts indicate organic (org) or bound forms of the element.
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F. Stuart Chapin III Pamela A. Mats
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Preface Human activities are affect
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Contents Preface . . . . . . . . .
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Contents ix Changes in Storage . .
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Contents xi Root Uptake Properties
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Contents xiii Disturbance . . . . .
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1 The Ecosystem Concept Ecosystem e
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Figure 1.1. Examples of ecosystems
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Community ecology Population ecolog
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ing from biotically driven changes
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The pool sizes and rates of cycling
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and alters patterns of vegetation d
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Figure 1.5. Direct and indirect eff
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many aspects of ecology, hydrology,
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Energy (W m -2 ) 1 1 0 1 0 1 0 1 Ab
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The Atmospheric System Atmospheric
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ecular O2 to form O3. The absorptio
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Hadley cell Hadley cell Ferrell cel
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early sailors as the doldrums. Subs
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phere, extends to depths of 75 to 2
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tures in Great Britain and western
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when the water vapor condenses to f
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Solar flux 1.4 1.2 1.0 0.8 0.6 0.4
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Figure 2.16. Time course of the ave
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Radiative forcing (W m -2 ) Warming
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January September Sun March pattern
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access to light compete effectively
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the top? How does each of these atm
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(Dokuchaev 1879, Jenny 1941, Amunds
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Organic carbon (%) Erosion likely g
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erosion dominating on steep hillslo
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conductivity of soils. Groundwater
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chemical weathering through their c
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Although most of the transfers in s
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leached material from above, it is
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opment, so these soils have weak de
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y roots and bacteria are important
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Table 3.4. Sequence of H + - consum
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new chemical conditions cause them
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72 4. Terrestrial Water and Energy
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98 5. Carbon Input to Terrestrial E
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100 5. Carbon Input to Terrestrial
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124 6. Terrestrial Production Proce
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152 7. Terrestrial Decomposition So
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154 7. Terrestrial Decomposition su
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156 7. Terrestrial Decomposition a
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158 7. Terrestrial Decomposition Ma
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160 7. Terrestrial Decomposition Re
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162 7. Terrestrial Decomposition cy
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Page 183 and 184: 8 Terrestrial Plant Nutrient Use In
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Page 205 and 206: 198 9. Terrestrial Nutrient Cycling
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Page 231 and 232: 10 Aquatic Carbon and Nutrient Cycl
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Page 251 and 252: 11 Trophic Dynamics Introduction Al
Page 253 and 254: 246 11. Trophic Dynamics People, fo
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Page 265 and 266: 258 11. Trophic Dynamics 4 th 3 rd
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Page 269 and 270: 262 11. Trophic Dynamics R R trophi
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266 12. Community Effects on Ecosys
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282 13. Temporal Dynamics An emergi
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284 13. Temporal Dynamics Small rod
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286 13. Temporal Dynamics regime (H
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288 13. Temporal Dynamics successio
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290 13. Temporal Dynamics Stage Lif
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292 13. Temporal Dynamics that redu
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294 13. Temporal Dynamics Soil carb
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296 13. Temporal Dynamics Nutrient
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298 13. Temporal Dynamics are aband
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300 13. Temporal Dynamics leaf area
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302 13. Temporal Dynamics account f
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304 13. Temporal Dynamics 6. How do
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306 14. Landscape Heterogeneity and
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15 Global Biogeochemical Cycles The
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tion of surface waters. On daily to
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Crowley 1995). An improved understa
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therefore limits the long-term rate
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important for understanding the rec
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Terrestrial N fixation (Tg yr -1 )
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The addition of limiting nutrients
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has a significant atmospheric compo
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cled. Evaporation and precipitation
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Figure 15.11. Trends in (A) world p
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which cycles are soil pools and flu
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Our use, mismanagement, and uninten
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mycorrhizal or nitrogen-fixing mutu
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Major human impacts on the natural
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promote long-term sustainability of
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Coral reef ecosystems have recently
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Table 16.3. Examples of services pr
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anthropogenic change and to sustain
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Abbreviations an nutrient productiv
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Abbreviations 373 NEE net ecosystem
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Glossary A horizon. Uppermost miner
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Biomass. Quantity of living materia
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Coriolis effect. Tendency, due to E
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Exoenzyme. Enzyme that is secreted
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Inceptisol. Soil order characterize
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Microbial loop. Microbial food web
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Phototroph. Nitrogen-fixing microor
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simply to the greater number of spe
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Sun leaf. Leaf that is acclimated t
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Color Plate I monthly averages of t
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Plate 3. The global pattern of net
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