List of boxes ................................................ xiii
Preface ........................................................ xv
Prologue
Environmental changes and ecosystem effects: two historical
examples ........................................................ 1
Acid rain ....................................................... 1
Global warming .................................................. 4
Approaches to solutions ......................................... 5
Further reading ................................................. 6
SECTION I HISTORY AND CONCEPTS .................................. 7
1 History of ecology ........................................... 8
Protoecology ................................................. 8
Early ecology ................................................ 9
fee ecosystem concept ....................................... 11
Modern ecology .............................................. 12
Biome ecology ............................................... 13
Alternative biome ecology: the Hubbard Brook project ........ 14
Postbiome ecosystem research ................................ 14
Later ecosystem research .................................... 15
Ecosystem research requires long-term observations and
aspects ..................................................... 16
Landscape ecology ........................................... 16
Gaia and non-equilibrium ecology ............................ 16
Sustainability and biodiversity ............................. 17
Further reading ............................................. 17
2 I Ecology, ecosystem and ecosystem science .................. 18
Ecology ..................................................... 18
Ecosystem ................................................... 20
Ecosystem science or ecosystem ecology? ..................... 21
Further reading ............................................. 24
3 Ecosystem ecology: cornerstones and scientific
methodology ................................................. 25
A note on terminology ....................................... 25
Understanding processes ..................................... 25
Basic principles and scientific methodology ................. 29
Cornerstones ................................................ 30
Mass balance ............................................. 30
Steady state ............................................. 31
Limiting nutrients ....................................... 32
Optimality ............................................... 32
One equation says more than a thousand words ................ 34
Understanding the cornerstones .............................. 35
Further reading ............................................. 38
SECTION II ECOSYSTEM STRUCTURE 39 AND FUNCTION
4 I Ecosystem structure: site factors, soil and vegetation .... 40
Terrestrial ecosystems and site factors ..................... 40
Soil physics and chemistry .................................. 41
The soil in an ecosystem context ......................... 41
Soil physical properties ................................. 42
Soil air ................................................. 45
Soil water ............................................... 46
Soil chemical properties, mineral nutrients and plants ... 46
Soil reactions and availability of mineral nutrients ..... 47
Soil types ............................................... 49
Ecosystem layering .......................................... 51
Terrestrial biomes .......................................... 55
Vegetation ............................................... 55
Features of terrestrial biomes and ecosystems ............ 56
Further reading ............................................. 62
5 Energy and water ............................................ 64
Solar energy ................................................ 64
Water balance ............................................... 72
Evapotranspiration .......................................... 75
Potential and actual evapotranspiration ..................... 78
SPAC: The soil-plant-atmosphere continuum ................... 82
Further reading ............................................. 87
6 Plant production ............................................ 88
Photosynthesis at leaf level ................................ 88
C3, C4 and CAM photosynthesis ............................... 92
Leaf area index and specific leaf area ...................... 95
Light extinction within a canopy ............................ 98
Photosynthesis at canopy level ............................. 102
Light-use efficiency ....................................... 103
Water-use efficiency ....................................... 105
Plant respiration .......................................... 106
Growth respiration ...................................... 106
Maintenance respiration ................................. 108
Root respiration and ion uptake ......................... 108
Plant nutrient relationships ............................... 108
Nutrient productivity ................................... 109
Nutrient-use efficiency ................................. 111
Plant allocation ........................................... 112
Plant nutrient uptake ...................................... 117
Steady-state nutrition ..................................... 121
Global plant traits ........................................ 121
Further reading ............................................ 123
7 Soil organic matter dynamics ............................... 124
Litter and soil organic matter ............................. 124
A model of litter decomposition ............................ 125
Litter quality ............................................. 130
Abiotic controls ........................................... 135
Extracellular enzymes ...................................... 137
Other controlling factors .................................. 139
Element concentration in decomposers ....................... 139
Soil food webs ............................................. 140
Models of soil organic matter .............................. 141
Further reading ............................................ 144
8 Organisms and ecosystem processes .......................... 145
Species and ecosystems ..................................... 145
Concepts of stability ...................................... 145
Species matter ............................................. 147
Or maybe not ............................................... 149
Further reading ............................................ 150
9 Element cycles ............................................. 151
Ecosystem scale ............................................ 153
The carbon cycle ........................................ 153
The nitrogen cycle ...................................... 161
The phosphorus cycle .................................... 169
The potassium cycle ..................................... 171
The calcium cycle ....................................... 172
The magnesium cycle ..................................... 173
The sulfur cycle ........................................ 174
Element cycling in a temperate Norway spruce forest ..... 175
Comparison of element cycling in different terrestrial
ecosystems .............................................. 180
Biomass .............................................. 180
Production ........................................... 180
Plant biomass turnover ............................... 182
Nitrogen-use efficiency: NUE ......................... 183
Degree of openness ................................... 184
Turnover time of soil carbon ......................... 185
Turnover time of soil nitrogen ....................... 185
Soil carbon/plant carbon ratio ....................... 186
Soil nitrogen/plant nitrogen ratio ................... 186
Global scale ............................................... 186
The carbon cycle ........................................ 186
The nitrogen cycle ...................................... 188
The phosphorus cycle .................................... 190
The sulfur cycle ........................................ 191
Human influence on element mass balances ................... 191
Further reading ............................................ 198
10 Principles ................................................. 200
Principles pertaining to boundary conditions (B) ........... 200
Principle Bl. On boundaries and storage ................. 200
Principle B2. On perturbing boundaries .................. 202
Principle B3. On nitrogen vs. phosphorus limitation ..... 202
Principle B4. On production and openness ................ 203
Principle B5. On time scales and openness ............... 203
Principles pertaining to energy and water (A) .............. 204
Principle AI. On climatic and ecosystem distribution .... 204
Principles pertaining to plant processes (P) ............... 204
Principle PI. On element availability ................... 204
Principle P2. On light limitation ....................... 206
Principle P3. On nutrient limitation .................... 206
Principle P4. On water limitation ....................... 206
Principle P5. On resource use efficiency and
acquisition ............................... 206
Principles pertaining to soil processes (S) ................ 207
Principle SI. On decomposition - energy limitation ...... 207
Principle S2. On nitrogen fixation ...................... 208
Principle S3. On nitrogen leaching ...................... 208
Principles pertaining to element cycles (E) ................ 208
Principle El. On openness of element cycles ............. 208
Principle E2. On element distributions .................. 209
Principle E3. On ecosystem carbon storage ............... 210
Further reading ............................................ 212
SECTION III ECOSYSTEM DYNAMICS AT DIFFERENT TIME SCALES ....... 213
11 Tectonic to orbital changes ................................ 215
Changes during the development of the biosphere ............ 215
Further reading ............................................ 224
12 Millennial to centennial or postglacial changes ............ 225
Postglacial development .................................... 225
Humans, climate and ecosystems ............................. 231
Centennial to recent changes ............................... 232
13 Centennial to annual changes ............................... 235
Wind ....................................................... 235
Fire ....................................................... 236
Grazing and other animal impacts ........................... 238
The tropical case ....................................... 238
The boreal case ......................................... 241
Stages in ecosystem development ............................ 242
Further reading ............................................ 244
SECTION IV APPLICATIONS ....................................... 245
14 Air pollution and forest ecosystems ........................ 246
Direct effects of air pollution on trees ................... 248
Indirect effects of air pollution on forest ecosystems ..... 250
Empirical evidence ...................................... 250
Forest soil acidification: centennial to recent
changes .............................................. 250
Biological versus pollution-generated acidification:
the hydrogen ion budget approach ..................... 253
Nutritional changes in trees ......................... 254
Growth rate changes in forests ....................... 255
Vegetation changes ................................... 257
Experimental evidence: understanding ....................... 258
Critical loads and the future .............................. 260
Critical loads from a European perspective ................. 261
Further reading ............................................ 263
15 Global change .............................................. 264
Effects on ecosystem carbon balance ........................ 265
Atmospheric carbon dioxide .............................. 265
Temperature ............................................. 266
Nitrogen ................................................ 267
Precipitation ........................................... 267
Effects on ecosystem nitrogen balance ...................... 268
Interactions ............................................... 269
Threshold effects .......................................... 270
Bioenergy: cure or curse? .................................. 271
Further reading ............................................ 274
Epilogue I Society and terrestrial ecosystem ecology ....... 275
What are ecosystem services? ............................... 276
Ecosystem changes in the last 50 years ..................... 277
Linking society and the scientific community ............... 280
Further reading ............................................ 283
Appendix 1 Abbreviations ...................................... 284
Appendix 2 Glossary ........................................... 287
Appendix 3 Some useful values and symbols used to represent
them ....................................................... 295
Appendix 4 Information and data on selected ecosystems ........ 296
(1) Arctic tundra: Stordalen, Sweden ....................... 296
Further reading ........................................ 297
(2) Boreal coniferous pine forest: Jдdraдs, Sweden ......... 297
Further reading ........................................ 297
(3) Temperate short grass prairie (steppe): Pawnee site,
Colorado, USA .......................................... 297
Further reading ........................................ 298
(4) Temperate beech forest: Solling {Heinz Ellenberg),
Germany ................................................ 298
Further reading ........................................ 298
(5) Temperate planted spruce forest: Skogaby, Sweden ....... 298
Further reading ........................................ 299
(6) Temperate coniferous rain forest: H.J. Andrews,
Oregon, USA ............................................ 299
Further reading ........................................ 299
(7) Tropical savanna: Point Noire (Kondi), Congo ........... 299
Further reading ........................................ 300
(8) Tropical dry forest: Chamela, Mexico ................... 300
Further reading ........................................ 300
(9) Tropical rain forest: San Carlos, Venezuela ............ 301
Further reading ........................................ 301
References .................................................... 304
Index ......................................................... 320
Colour plate section is found between pp. 174 and 175.
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