The Ecology of Phytoplankton (Ecology, Biodiversity and Conservation)

Cover......Page 1
Half-title......Page 3
Series title......Page 4
Title......Page 5
Copyright......Page 6
Dedication......Page 7
Contents......Page 9
Preface......Page 13
Acknowledgements......Page 16
1.1 Definitions and terminology......Page 17
1.2 Historical context of phytoplankton studies......Page 19
1.3 The diversification of phytoplankton......Page 20
1.4 General features of phytoplankton......Page 31
1.4.2 Regulating surface-to-volume ratio......Page 35
1.5 The construction and composition of freshwater phytoplankton......Page 40
1.5.1 Dry weight......Page 41
1.5.2 Skeletal silica......Page 43
1.5.3 Organic composition......Page 44
1.5.4 Chlorophyll content......Page 49
1.6 Marine phytoplankton......Page 50
1.7 Summary......Page 52
2.1 Introduction......Page 54
2.2.1 Physical properties of water......Page 55
2.2.2 Generating oceanic circulation......Page 57
2.3.1 Generation of turbulence......Page 58
2.3.2 Quantifying turbulence......Page 62
2.3.3 Turbulent dissipation......Page 63
2.3.4 Turbulent embedding of phytoplankton......Page 64
2.4 Phytoplankton sinking and floating......Page 65
2.4.2 Departure from spherical shape: form resistance......Page 66
2.5.2 Density......Page 69
Lipid accumulation......Page 70
Mucilage......Page 71
Gas vacuoles......Page 72
Chain formation......Page 76
Cylinder elongation and cylindrical filaments......Page 78
Colony formation......Page 79
2.5.4 Vital regulation of sinking rate......Page 81
2.6.1 Sinking, floating and entrainment......Page 83
2.6.2 Loss of sinking particles from turbulent layers......Page 86
2.6.3 Mixed depth variability in natural water columns......Page 87
2.6.4 Vertical structure in the pelagic......Page 89
2.6.5 Mixing times......Page 91
2.6.6 Particle settling from variable mixed layers: an experiment......Page 92
2.7 The spatial distribution of phytoplankton......Page 93
2.7.1 Vertical distribution of phytoplankton......Page 95
Non-motile, negatively buoyant plankters......Page 96
Positively buoyant plankters......Page 97
Motile plankters......Page 98
Small-scale patchiness......Page 100
Patchiness in small lake basins......Page 102
Large-scale patchiness......Page 104
Relevance of patchiness......Page 105
2.8 Summary......Page 106
3.1 Introduction......Page 109
3.2 Essential biochemistry of photosynthesis......Page 110
3.2.1 Light harvesting, excitation and electron capture......Page 111
3.2.2 Photosystem architecture......Page 112
3.2.3 Carbon reduction and allocation......Page 114
3.3.1 Measurement of light-dependent photosynthetic oxygen production......Page 117
3.3.2 Measurement of light-dependent photosynthetic carbon fixation......Page 121
Characterising the photosynthetic impact of the underwater light intensity......Page 124
The impact of optical properties of water on the underwater light spectrum......Page 126
Phytoplankton adaptations to sub-saturating irradiances......Page 129
Photosynthetic limits in lakes and seas......Page 132
Photoadaptation to vertical mixing......Page 133
3.3.4 Photoprotection, photoinhibition and photooxidation......Page 137
Avoidance reactions......Page 138
Excretion......Page 139
3.4.1 Sources and fluxes of available inorganic carbon......Page 140
3.4.2 Phytoplankton uptake of carbon dioxide......Page 143
3.4.3 Species-specific sensitivities to low DIC concentrations......Page 144
3.5 Capacity, achievement and fate of primary production at the ecosystem scale......Page 147
3.5.1 Primary production at the local scale......Page 148
3.5.2 Primary production at the global scale......Page 150
3.5.3 Capacity......Page 152
3.5.4 Photosynthetic yield to the planktic food web......Page 155
3.6 Summary......Page 159
4.1 Introduction......Page 161
4.2.1 Supply of nutrients......Page 162
4.2.2 Moving nutrients into the cell......Page 164
4.2.3 Empirical models of nutrient uptake......Page 166
4.3.1 The sources and biological availability of phosphorus in natural waters......Page 167
4.3.2 MRP-uptake kinetics......Page 171
4.3.3 Metabolic-rate limitation by phosphorus......Page 174
4.3.4 Capacity limitation and potential phosphorus yield......Page 175
4.4 Nitrogen: requirements, sources, uptake and metabolism in phytoplankton......Page 177
4.4.1 The sources and availability of nitrogen to phytoplankton......Page 178
4.4.2 Uptake of DIN by phytoplankton......Page 179
4.4.3 Nitrogen fixation......Page 180
4.5 The role of micronutrients......Page 182
4.5.1 The toxic metals......Page 183
4.5.2 Iron......Page 184
4.5.3 Organic micronutrients and vitamins......Page 186
4.6.1 Cations......Page 187
4.6.2 Anions......Page 188
4.7 Silicon: requirements, uptake, deployment in phytoplankton......Page 189
4.8 Summary......Page 191
5.1 Introduction: characterising growth......Page 194
5.2.1 The cell growth cycle......Page 195
5.2.2 Cell division and population growth......Page 198
5.3.1 Maximum replication rates as a function of algal morphology......Page 199
5.3.2 The effect of temperature......Page 202
5.3.3 Resourcing maximal replication......Page 204
5.4.1 The effect of truncated photoperiod......Page 205
5.4.2 The effect of persistent low light intensities......Page 208
5.4.3 The effect of fluctuating light......Page 209
Phosphorus deficiency......Page 210
Nitrogen deficiency......Page 211
Silicon deficiency......Page 212
Resource-based competition......Page 213
Stoichiometry......Page 215
Chronic nutrient deficiencies......Page 219
Discontinuous nutrient deficiences......Page 220
Low insolation and growth-rate regulation......Page 222
Trait interaction and functional differentiation in phytoplankton......Page 223
Growth and reproductive strategies......Page 224
5.4.6 Resource exhaustion and survival......Page 227
Growth rates from episodic events......Page 233
Frequency of cell division......Page 234
Growth from the depletion of resource......Page 236
5.5.2 The spring increase in temperate lakes: the case of Windermere......Page 237
5.5.3 Selection by performance......Page 241
5.5.4 Temporal changes in performance selection......Page 247
5.5.5 Modelling growth rates in field......Page 250
5.6 Summary......Page 252
6.1 Introduction......Page 255
6.2.1 Expressing dilution......Page 256
6.2.2 Dilution in the population ecology of phytoplankton......Page 257
6.2.3 Phytoplankton population dynamics in rivers......Page 258
6.3.1 Loss by sinking......Page 259
6.3.2 Mixed depth and the population dynamics of diatoms......Page 260
6.3.3 Accumulation and resuspension of deposited material......Page 265
6.4 Consumption by herbivores......Page 266
Protistan microzooplankton......Page 267
Multicellular microzooplankton......Page 275
Freshwater mesozooplankton......Page 276
Marine mesozooplankton......Page 277
Planktivorous macroplankton, megaplankton and nekton......Page 278
6.4.2 Impacts of filter-feeding on phytoplankton......Page 280
Filtration rates......Page 281
Food availability......Page 283
Algal removal and grazer nutrition......Page 286
Food thresholds and natural populations......Page 292
6.4.3 Selective feeding......Page 294
6.4.4 Losses to grazers......Page 296
6.4.5 Phytoplankton–zooplankton interactions......Page 301
Competitive interactions......Page 302
Bottom–up and top–down processes in oligotrophic systems......Page 303
Bottom–up and top–down processes in enriched systems......Page 304
Intervention in food-web interactions......Page 306
Food-chain length......Page 307
6.5.1 Fungal parasites......Page 308
6.5.2 Protozoan and other parasites......Page 310
6.5.3 Pathogenic bacteria and viruses......Page 311
6.6 Death and decomposition......Page 312
6.7 Aggregated impacts of loss processes on phytoplankton composition......Page 313
6.8 Summary......Page 316
7.2.1 Species composition in the sea......Page 318
Phytoplankton of the North Pacific Subtropical Gyre......Page 320
Phytoplankton of deep boreal waters......Page 322
Phytoplankton of the Southern Ocean......Page 323
Upwellings......Page 324
Shelf phytoplankton......Page 325
Coastal and near-shore waters......Page 326
7.2.2 Species assemblage patterns in the sea......Page 329
7.2.3 Species assemblage patterns in lakes......Page 334
Phytoplankton of large oligotrophic and ultraoligotrophic lakes......Page 335
Phytoplankton of smaller temperate oligotrophic lakes......Page 341
Phytoplankton of sub-Arctic lakes......Page 344
Phytoplankton in selected regions: medium-sized glacial lakes of the European Alps......Page 345
Phytoplankton in selected regions: small......Page 346
Phytoplankton in selected regions: glacial lakes of Araucania......Page 351
Phytoplankton of small kataglacial lakes......Page 352
Phytoplankton of large, low-latitude lakes......Page 355
Phytoplankton in shallow lakes......Page 358
7.2.4 Species assemblage patterns in lakes......Page 361
7.3.1 Assembly of nascent communities......Page 366
Minimal communities......Page 367
The pioneer element of nascent communities......Page 368
Species richness of nascent communities......Page 370
Ascendency: power and exergy......Page 371
Ascendency: environmental constraints......Page 373
Succession......Page 375
Filtration and community assembly......Page 376
7.3.3 Species diversity and disequilibrium in natural communities......Page 383
Disequilibrium......Page 385
Resource competition as a mechanism of coexistence......Page 387
Disturbance as a mechanism of coexistence......Page 388
IDH and diversity......Page 390
Explaining the plankton paradox: where does the diversity come from?......Page 395
7.3.4 Stability and fidelity in phytoplankton communities......Page 397
7.3.5 Structure and dynamics of phytoplankton community assembly......Page 399
7.4 Summary......Page 401
8.2.1 Fate of primary product in the open pelagic......Page 403
8.2.2 Food and recruitment of consumers in relation to primary production......Page 404
8.2.3 Carbon subsidies in lakes and the sea......Page 406
8.2.4 The relationship between energy flow and structure in the plankton......Page 409
8.3 Anthropogenic change in pelagic environments......Page 411
Defining the issues......Page 413
Aquatic impacts of phosphorus loads to lakes......Page 415
Cyanobacterial ‘blooms’......Page 417
Cyanobacterial toxicity......Page 418
Control of cyanobacteria......Page 421
Red tides......Page 423
8.3.3 Controlling eutrophication by phosphorus load reduction......Page 424
8.3.4 The internal load problem......Page 428
8.3.5 Physical methods to control phytoplankton abundance......Page 430
8.3.6 The control of phytoplankton by biological manipulation......Page 432
Nutrients, phytoplankton and grazers......Page 433
The role of macrophytes......Page 434
Effects of eutrophication on macrophytes......Page 436
Macrophytes and trophic relationships......Page 437
Biomanipulative management......Page 438
8.3.7 Phytoplankton and acidification......Page 439
8.3.8 Anthropogenic effects on the sea......Page 441
Anthropogenic impacts upon marine communities and food webs......Page 442
Anthropogenic countermeasures to the atmospheric accumulation of greenhouse gases......Page 444
Phytoplankton and the future......Page 447
8.4 Summary......Page 448
8.5 A last word......Page 451
Glossary......Page 453
Symbols......Page 456
Variables......Page 457
Abbreviations used in the text......Page 462
References......Page 463
LAKES AND RESERVOIRS......Page 524
Index to genera and species of phytoplankton......Page 527
Index to genera and species of organisms
other than phytoplankton......Page 536
General index......Page 540