Colin S. Reynolds VEGETATION PROCESSES IN THE PELAGIC: A MODEL FOR ECOSYSTEM THEORY Introduction (Otto Kinne) Colin S. Reynolds: A Laudatio (William D. Williams) Publisher: Ecology Institute Nordbunte 23, D-21385 Oldendorf/Luhe Germany
Contents Introduction (Otto Kinne) Colin S. Reynolds: A Laudatio (William D. Williams) Dankwort Preface XI XXI XXIII XXIII I II THE PELAGIC VEGETATION (1) Introduction: the phytoplankton 1 (2) Marine phytoplankton associations 4 (3) Freshwater phytoplankton associations 6 (4) Diatom-dominated plankton 7 (5) Chrysophycean plankton 10 (6) Oligotrophic chlorococcal plankton 11 (7) Eutrophic chlorophyte plankton 12 (8) Dinoflagellate plankton 12 (9) Cyanobacteria-dominated plankton 13 (10) Cryptomonad plankton 15 (11) Nanoplankton 16 (12) Picoplankton 16 (13) Photosynthetic bacterioplankton 17 (14) Miscellaneous categories 17 (15) Explaining the occurrence of phytoplankton 18 Summary 19 THE PELAGIC ENVIRONMENT (1) Introduction 21 (2) Critical physical properties of water 23 (3) Turbulence 24 (4) Turbulent dissipation 30 (5 ) Turbulent embedding of phytoplankton 31 (6) Sinking and floating 33 (7) Entrainment and disentrainment 35 (8) Turbulent extent 36 (9) Mixing times 46 (10) Light reception in mixed layers 48
(11) Factors controlling the chemical composition of pelagic environments 51 (12) Nutrients in pelagic environments 53 (13) Nitrogen 55 (14) Phosphorus 56 (15) Carbon dioxide 63 (16) Oxygen and redox 64 (17) Metabolic sensitivity of lakes to nutrient loads 67 (18) Silicon and iron 68 Summary. 70 III IV RESOURCE GATHERING IN PELAGIC PLANTS (1) Introduction 73 (2) Overview of cellular uptake and intracellular transport 74 (3) Resource gathering at the molecular level 78 (4) The molecular basis of photosynthetic carbon fixation 79 (5) Light-dependent photosynthetic rates 82 (6) Dealing with super-saturating photon fluxes 85 (7) Photoadaptation to low light intensities 89 (8) Carbon uptake by pelagic plants 91 (9) Phosphorus 94 (10) Nitrogen 97 (11) Starvation and survival 98 Summary 100 GROWTH OF PELAGIC PLANTS (1) Introduction 101 (2) Thecellgrowthcycle 102 (3) Cell division 105 (4) Replication rates under ideal conditions 107 (5) The effect of temperature on replication rates ' 111 (6) Resourcing maximal replication. 113 (7) Replication rates under sub-ideal conditions: the effect of photoperiod 114 (8) Replication rates under sub-ideal conditions: the effect of persistent low light intensities 120 (9) Replication rates under sub-ideal conditions: the effect of nutrient deficiency 121 (10) Resource ratios 122 (11) Functional differentiation of phytoplankton 125 Summary 126
V VI DEVELOPMENT OF PELAGIC PLANT POPULATIONS (1) Introduction 129 (2) Pelagic biomass capacities 131 (3) Supportive capacities and lake typology 136 (4) Filling the capacity: in situ replication rates 142 (5) Replication rates in energy-deficient environments 147 (6) Replication in chronically resource-deficient environments 152 (7) Replication in discontinuously resource-deficient environments 154 (8) Perennation of populations 158 (9) Replication strategies 161 (10) The selection of natural pelagic populations 167 Summary 171 DEPLETION OF PELAGIC PLANT POPULATIONS (1) Introduction 173 (2) Wash-out and dilution 174 (3) Sinking and sedimentation 177 (4) Mixed depth and phytoplankton dynamics: regulating sinking rate 180 (5) Mixed depth and the population dynamics of diatoms 184 (6) Consumption by heterotrophs 188 (7) Filter feeding 191 (8) Selective feeding 195 (9) Microbial food pathways 196 (10) Pathogens 197 (11) Death and decomposition 198 (12) Loss rates and the selection of phytoplankton 199 Summary 201 VII ASSEMBLY OF PELAGIC PLANT COMMUNITIES (1) Introduction 205 (2) Organisation of the true pelagic 206 (3) Organisation and opportunities in small lakes 209 (4) Alternative vegetation states in small and shallow lakes 211 (5) Organisation and opportunities in intermediate-sized lakes 213 (6) Community assembly and the habitat template 215 (7) Temporal changes in community structure 222 (8) Ecological succession 224 (9) Succession in pelagic communities 227 (10) Climax in pelagic successions. 231
(11) Experiments in pelagic succession 232 (12) Species diversity through successions 236 (13) Disturbances to pelagic successions 240 (14) Severity and frequency of disturbances 242 (15) Plankton and the Intermediate Disturbance Hypothesis 248 (16) Interannual variability in community structure 252 Summary 256 VIII REGULATION OF PELAGIC ECOSYSTEMS (1) Introduction 259 (2) The stability of pelagic ecosystems 259 (3) Pulsed resource exploitation in pelagic ecosystems 263 (4) Bottom-up or top-down control? 265 (5) The energetic control of pelagic ecosystems 270 (6) The energetic state of pelagic ecosystems 274 (7) Exergy and pelagic heterogeneity 277 (8) Structural classification of pelagic plant communities 281 (9) Pelagic plagioclimaces 284 (10) Pelagic ecosystem regulation 286 Summary 289 IX LESSONS AND APPLICATIONS (1) Introduction 291 (2) Harvesting sunlight 291 (3) Ecosystem variability 293 (4) Ecosystem exploitation 295 (5) Whither biodiversity? 298 (6) Water resources: quantity and quality 301 (7) Eutrophication 304 (8) Correcting eutrophication: nutrient-load reduction 306 (9) Controlling phytoplankton biomass: physical mixing 311 (10) Controlling phytoplankton biomass: biomanipulation 312 (11) The problem of algal toxicity 316 (12) Treating acidified lakes 320 (13) Sustainable management of water resources 322 (14) The ecosystems approach to water management 325 (15) Tailpiece 327 Summary 327 Glossary of symbols 331 References 339