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Contents Preface xvii Copyright acknowledgements xix 1 Microbial diversity and freshwater ecosystems 1 1.1 General introduction 1 1.1.1 The aquatic existence 1 1.1.2 The global water supply - limnology and oceanography 1 1.1.3 Freshwater systems: some terms and definitions 3 1.1.4 The biology of freshwater microorganisms 4 A. BIOLOGICAL DIVERSITY IN THE FRESHWATER ENVIRONMENT 4 1.2 Biodiversity of microorganisms 4 1.2.1 Domains of life 4 1.2.2 Size range 6 1.2.3 Autotrophs and heterotrophs 7 1.2.4 Planktonic and benthic microorganisms 10 1.2.5 Metabolically active and inactive states 11 1.2.6 Evolutionary strategies: r-selected and K-selected organisms 12 1.3 Biodiversity in ecosystems, communities, and species populations 15 1.3.1 Main ecosystems 15 1.3.2 Diversity within subsidiary communities 16 1.3.3 Biodiversity within single-species populations 16 B. ECOSYSTEMS 17 1.4 The biofilm community: a small-scale freshwater ecosystem 18 1.4.1 Interactions between microorganisms 19 1.4.2 Biomass formation and transfer 20 1.4.3 Maintenance of the internal environment 20 1.4.4 Interactions with the external environment 21 1.5 The pelagic ecosystem: a large-scale unit within the lake environment 21 1.5.1 Interactions between organisms 21 1.5.2 Trophic connections and biomass transfer 23 1.5.3 Maintenance of the internal environment 28 1.5.4 Interactions with the external environment 28 1.6 Homeostasis and ecosystem stability 29 1.6.1 Stress factors 30 1.6.2 General theoretical predictions: the community response 30 1.6.3 Observed stress responses: from molecules to communities 31 1.6.4 Assessment of ecosystem stability 31 1.6.5 Ecosystem stability and community structure 32 1.6.6 Biological response signatures 34 C. FOOD WEBS IN LENTIC AND LOTIC SYSTEMS 34 1.7 Pelagic food webs 34 1.7.1 Case study: microbial food web associated with an algal bloom 34 1.7.2 Case study: general food web in the water column of Lake Baikal (Russia) 36 1.8 Communities and food webs of running waters 40 1.8.1 Allochthonous carbon: dissolved and particulate matter in river systems 40 1.8.2 Pelagic and benthic communities 42 1.8.3 The microbial food web 43 2 Freshwater environments: the influence of physico-chemical conditions on microbial communities 47 A. INTRODUCTION 47 2.1 The aquatic medium: water, dissolved and particulate components 47 2.1.1 Particulate matter 47 2.1.2 Aquatic matrix 48 2.2 Freshwater environments 52 B. LAKES 52 2.3 Lake morphology and hydrology 53 2.3.1 Lake morphology 53 2.3.2 Lake hydrology and the surrounding terrestrial environment 57 2.4 Lakes as isolated environments 60 2.4.1 Isolated development 60 2.4.2 Lake Baikal: an ancient lake with a diverse and unique fauna and flora 60 2.5 Climatic influences on lakes 62 2.5.1 Temperate lakes - seasonal variations and lake stratification 63 2.5.2 Biological significance of stratification 65 2.5.3 Tropical lakes 66 2.5.4 Polar and sub-polar lakes 67 C. WETLANDS 68 2.6 General characteristics 68 2.6.1 Wetland diversity and global scale 68 2.6.2 Unifying features of wetlands 68 2.6.3 The role of wetlands in energy and material flow 69 2.7 Wetland habitats and communities 69 2.8 Case studies on wetland areas 70 2.8.1 Case study: Trebon basin biosphere reserve 71 D. STREAMS AND RIVERS 72 2.9 Comparison of lotic and lentic systems 72 2.10 River flow and the benthic community 73 2.10.1 Flow characteristics of lotic systems 73 2.10.2 Influence of water flow on benthic microorganisms 75 2.11 River hydrology 78 E. ESTUARIES 78 2.12 River inflow: water mixing, estuarine productivity, and eutrophication of coastal areas 79 2.12.1 Mixing of fresh and saltwaters 80 2.12.2 High productivity of estuarine systems 80 2.12.3 Eutrophication of surrounding coastal areas 81 2.13 Habitats and communities 82 2.13.1 Pelagic zone 82 2.13.2 Sediments and mudflats 82 F. ADVERSE AND EXTREME CONDITIONS IN FRESHWATER ENVIRONMENTS 84 2.14 Adverse conditions as part of the environmental continuum 85 2.14.1 Variations in oxygen concentration 85 2.14.2 Nutrient availability 85 2.14.3 Solar radiation 87 2.15 Extreme environmental conditions 87 2.15.1 Temperature 87 2.15.2 pH 89 2.15.3 Conditions of low water availability: saline environments 91 2.15.4 Conditions of low water availability: ice and snow environments 92 2.15.5 Variations in hydrostatic pressure 92 2.15.6 Organic and inorganic pollution 93 2.16 A potentially extreme microenvironment: the air-water surface 94 2.16.1 Chemical composition of the surface microlayer 95 2.16.2 Physical processes and transformations in the surface biofilm 96 2.16.3 Microbial community at the air-water interface 98 2.17 Microbial communities of snow and ice: life in the frozen state 98 2.17.1 Snow and ice as an extreme environment 99 2.17.2 Requirement for water in the liquid state 99 2.17.3 Snow ecosystems 99 2.17.4 The physical properties of snow 100 2.17.5 Snow and ice microorganisms 102 3 Algae: the major microbial biomass in freshwater systems 105 A. TAXONOMIC AND MOLECULAR CHARACTERIZATION 107 3.1 Major taxonomic divisions of freshwater algae 107 3.1.1 Microscopical appearance, motility and ecological features 107 3.1.2 Biochemical and cytological characteristics 110 3.1.3 General summary of the different groups 111 3.2 Algal species: taxonomy and intraspecific variation 114 3.2.1 Taxonomy of algal species 114 3.2.2 Chemical diversity within species - enzyme analysis, molecular groups, and elemental composition 114 3.3 Molecular analysis 116 3.3.1 Molecular characterization and identification of algae 116 3.3.2 Investigation of gene function in freshwater algae 119 B. SIZE, SHAPE, AND SURFACE MUCILAGE 122 3.4 Phytoplankton size and shape 122 3.4.1 Cell counts and biovolume 123 3.4.2 From picoplankton to macroplankton 123 3.4.3 Biological significance of size and shape 124 3.4.4 Variation in size and shape within phytoplankton populations 128 3.5 Mucilaginous and non-mucilaginous algae 130 3.5.1 Chemical composition of mucilage 131 3.5.2 Role of mucilage in phytoplankton activities 131 3.5.3 Environmental impact and biogeochemical cycles 133 C. ACTIVITIES WITHIN THE FRESHWATER ENVIRONMENT 133 3.6 Benthic algae: interactions with planktonic algae and ecological significance 133 3.6.1 Planktonic and benthic algae 133 3.6.2 Lake periphyton 136 3.6.3 Benthic algae in flowing waters 138 3.6.4 Ecological role of benthic algae 138 3.7 Temporal activities of freshwater algae 139 3.7.1 Short-term changes: molecular and cellular processes 140 3.7.2 Medium-term changes: algal succession 142 3.7.3 Long-term changes: variations over a number of years 146 3.8 Phytoplankton distribution within the water column 148 3.8.1 Case study: vertical zonation of phytoplankton in a stratified lake 148 3.8.2 Active migration of algae 149 3.8.3 Passive movement of algae within the water column 156 3.9 Freshwater algae and nutrient status of the environment 157 3.9.1 Phytoplankton species composition and lake nutrient status 157 3.9.2 Nutrient status of river environments - effect on benthic algal biofilms 160 D. STRATEGIES FOR SURVIVAL 161 3.10 Strategies for survival: the planktonic environment 161 3.10.1 Meroplanktonic algae 162 3.10.2 Strategies for unstable and stable environments: r-selected and K-selected algae 164 3.11 Heterotrophic nutrition in freshwater algae 165 3.11.1 Organotrophy 167 3.11.2 Phagotrophy 168 3.12 Survival in snow and ice: adaptations of cryophilic algae 171 3.12.1 Major groups of cryophilic algae 171 3.12.2 Life cycles of snow algae 172 3.12.3 Physiological adaptations of snow algae 174 E. BIODIVERSITY IN THE ALGAL COMMUNITY 177 3.13 Variety of freshwater algae: indices of species diversity 177 3.13.1 The paradox of phytoplankton diversity 177 3.13.2 Biodiversity indices 178 3.13.3 Numerical comparison of phytoplankton populations 179 3.13.4 Biodiversity and ecosystem function 180 4 Competition for light 183 4.1 The light environment 184 4.1.1 Physical properties of light: terms and units of measurement 184 4.1.2 Light thresholds for biological activities 185 4.1.3 Light under water: refraction, absorption, and scattering 186 4.1.4 Light energy conversion: from lake surface to algal biomass 188 4.2 Photosynthetic processes in the freshwater environment 190 4.2.1 Light and dark reactions 190 4.2.2 Photosynthetic microorganisms 191 4.2.3 Measurement of photosynthesis 191 4.2.4 Photosynthetic response to varying light intensity 192 4.3 Light as a growth resource 194 4.3.1 Strategies for light uptake and utilization 194 4.3.2 Light-photosynthetic response in different algae 195 4.3.3 Conservation of energy 196 4.3.4 Diversity in small molecular weight solutes and osmoregulation 197 4.4 Algal growth and productivity 198 4.4.1 Primary production: concepts and terms 198 4.4.2 Primary production and algal biomass 199 4.4.3 Field measurements of primary productivity 199 4.5 Photosynthetic bacteria 201 4.5.1 Major groups 202 4.5.2 Photosynthetic pigments 202 4.5.3 Bacterial primary productivity 203 4.6 Photoadaptation: responses of aquatic algae to limited supplies of light energy 204 4.6.1 Different aspects of light limitation 205 4.6.2 Variable light intensity: light-responsive gene expression 206 4.6.3 Photosynthetic responses to low light intensity 207 4.6.4 Spectral composition of light: changes in pigment composition 211 4.7 Carbon uptake and excretion by algal cells 212 4.7.1 Changes in environmental CO2 and pH 212 4.7.2 Excretion of dissolved organic carbon by phytoplankton cells 213 4.8 Competition for light and carbon dioxide between algae and higher plants 217 4.8.1 The balance between algae and macrophytes in different aquatic environments 217 4.8.2 Case study: competition between algae and macrophytes in shallow lakes of the Trebon wetlands 218 4.8.3 Physiological and environmental adaptations in the competition between algae and macrophytes 220 4.9 Damaging effects of high levels of solar radiation: photoinhibition 223 4.9.1 Specific mechanisms of photoinhibition 223 4.9.2 General effects of photoinhibition 226 4.9.3 Strategies for the avoidance of photoinhibition 227 4.9.4 Photoinhibition and cell size 229 4.9.5 Lack of photoinhibition in benthic communities 230 4.9.6 Photoinhibition in extreme high-light environments 230 4.10 Periodic effects of light on seasonal and diurnal activities of freshwater biota 232 4.10.1 Seasonal periodicity 232 4.10.2 Diurnal changes 233 4.10.3 Circadian rhythms in blue-green algae 234 4.10.4 Circadian rhythms in dinoflagellates 236 5 Inorganic nutrients: uptake and cycling in freshwater systems 237 5.1 Chemical composition of natural waters 237 5.1.1 Soluble inorganic matter in lakes and rivers 237 5.1.2 Aerial deposition of nutrients 239 5.1.3 Nutrient inflow from terrestrial sources 239 5.1.4 Chemical requirements and composition of freshwater biota 240 5.1.5 Case study: elemental composition of Ceratium hirundinella 242 5.1.6 Nutrient availability and cycling in aquatic systems 245 5.2 Nutrient uptake and growth kinetics 248 5.2.1 Empirical models for algal nutrient kinetics 248 5.2.2 Competition and growth in the aquatic environment 250 5.2.3 Nutrient availability and water movement 252 5.2.4 Acute nutrient deprivation as an environmental stress factor 253 A. NITROGEN 253 5.3 Biological availability of nitrogen in freshwater environments 253 5.3.1 Soluble nitrogenous compounds 253 5.4 The nitrogen cycle 256 5.4.1 Nitrate entry and uptake (soluble inorganic to insoluble organic nitrogen) 256 5.4.2 Complex organic nitrogen (biomass) transformations (successive states of insoluble organic nitrogen) 257 5.4.3 Remineralization (insoluble organic to soluble inorganic nitrogen) 257 5.4.4 Nitrification/denitrification (oxidation/reduction of soluble inorganic compounds) 257 5.5 Uptake of nitrate and ammonium ions by algae 259 5.5.1 Biochemical processes 259 5.5.2 Species variations in nitrate uptake 260 5.5.3 Environmental regulation of nitrate assimilation 260 5.5.4 Nitrogen uptake, CO2 assimilation, and photosynthesis 261 5.6 Nitrogen fixation 262 5.6.1 Ecological significance of nitrogen fixation 262 5.6.2 The nitrogenase enzyme and strategies of fixation 262 5.6.3 Heterocysts: nitrogen fixation by colonial blue-green algae 263 5.6.4 Diurnal control of nitrogen fixation: unicellular blue-green algae 264 5.6.5 Anaerobic environment: nitrogen-fixing bacteria 265 B. PHOSPHORUS 267 5.7 Occurrence and biological availability of phosphorus 267 5.7.1 Phosphorus availability and limitation 267 5.7.2 The phosphorus cycle 268 5.8 Adaptations of freshwater microorganisms to low phosphorus concentrations 271 5.8.1 Kinetics of phosphorus uptake 271 5.8.2 Luxury consumption of phosphate 271 5.8.3 Secretion of alkaline phosphatase 273 C. SILICON: A WIDELY-AVAILABLE ELEMENT OF LIMITED METABOLIC IMPORTANCE 274 5.9 The silicon cycle 274 5.10 Silicon and diatoms 276 5.10.1 Si uptake and phytoplankton succession 276 5.10.2 Si uptake and cell-wall formation 277 D. TRACE ELEMENTS 280 5.11 Biological role of trace elements 282 5.11.1 Environmental uptake of trace elements 282 5.11.2 Stimulation of growth in aquatic environments 282 5.11.3 Importance of trace metals in the culture of aquatic algae 283 5.11.4 Biochemical roles of trace elements 283 5.12 Cycling of iron and other trace metals in the aquatic environment 285 5.12.1 The iron cycle 285 5.12.2 The manganese cycle 288 6 Bacteria: the main heterotrophic microorganisms in freshwater systems 289 A. GENERAL DIVERSITY WITHIN THE ENVIRONMENT 289 6.1. General diversity, habitat preferences, and ecological significance of freshwater bacteria 289 6.1.1 General diversity 289 6.1.2 Habitat preferences 290 6.1.3 Environmental significance of freshwater bacteria 292 6.2 Taxonomic, biochemical, and molecular characterization of freshwater bacteria 293 6.2.1 Species identification 293 6.2.2 Genetic markers: detection of particular strains in the aquatic environment 294 6.2.3 Biochemical characterization of bacterial communities 295 6.2.4 Case study: changes in bacterial community function and composition as a response to variations in the supply of dissolved organic material (DOM) 295 B. GENETIC INTERACTIONS 296 6.3 Genetic diversity 296 6.3.1 Chromosomal and accessory DNA 296 6.3.2 The ecological importance of gene transfer in freshwater systems 297 6.3.3 Total genetic diversity: the 'community genome' 298 6.4 Mechanisms for gene transfer in freshwater systems 299 6.4.1 Transformation: uptake of exogenous DNA 299 6.4.2 Transduction: gene transfer between bacteria via bacteriophages 302 6.4.3 Conjugation: transfer of plasmid DNA by direct cell contact 302 6.5 Evidence for gene transfer in the aquatic environment 302 6.5.1 Retrospective analysis 302 6.5.2 Case study: plasmid-borne resistance in aquatic bacteria 303 6.5.3 Laboratory (in vitro) studies on plasmid transfer 303 6.5.4 Case study: plasmid transfer in Pseudomonas aeruginosa 304 6.5.5 Field (in situ) studies on bacterial gene transfer 305 C. METABOLIC ACTIVITIES 306 6.6 Metabolic diversity of freshwater bacteria 306 6.6.1 Key metabolic parameters 306 6.6.2 CO2 fixation 306 6.6.3 Breakdown of organic matter in aerobic and anaerobic environments 307 6.6.4 Bacterial adaptations to low-nutrient environments 312 6.7 Photosynthetic bacteria 314 6.7.1 General characteristics 314 6.7.2 Motility 314 6.7.3 Ecology 316 6.8 Bacteria and inorganic cycles 316 6.8.1 Bacterial metabolism and the sulphur cycle 317 D. BACTERIAL POPULATIONS AND PRODUCTIVITY 318 6.9 Bacterial populations 318 6.9.1 Techniques for counting bacterial populations 318 6.9.2 Biological significance of total and viable counts 319 6.10 Bacterial productivity 320 6.10.1 Measurement of productivity 320 6.10.2 Regulation of bacterial populations and biomass 321 6.10.3 Primary and secondary productivity: correlation between bacterial and algal populations 322 6.10.4 Primary and secondary productivity: the role of dissolved organic carbon 323 6.10.5 Bacterial productivity and aquatic food webs 325 E. BACTERIAL COMMUNITIES IN THE LOTIC ENVIRONMENT 326 6.11 Bacterial Biofilms 326 6.11.1 The development of biofilms 326 6.11.2 Dynamic interactions in the establishment of biofilms: the role of bacterial co-aggregation 328 6.11.3 Case study: specific recognition and adhesion amongst aquatic biofilm bacteria 328 F. BACTERIAL INTERACTIONS WITH PHYTOPLANKTON 330 6.12 Interactions between phytoplankton and planktonic bacteria 330 6.12.1 Competition for inorganic nutrients 330 6.12.2 Antagonistic interactions between bacteria and algae 331 6.13 Epiphytic associations of bacteria with phytoplankton 334 6.13.1 Bacteria within the phycosphere 335 6.13.2 Observation and enumeration of epiphytic bacteria 336 6.13.3 Specific associations between bacteria and blue-green algae 338 7 Viruses: major parasites in the freshwater environment 341 7.1 Viruses as freshwater biota 341 7.1.1 General role in the freshwater environment 341 7.1.2 Major groups and taxonomy of freshwater viruses 342 7.2 The virus life cycle: intracellular and free viral states 342 7.2.1 Significance of the lysogenic state 343 7.3 Detection and quantitation of freshwater viruses 344 7.3.1 Free particulate viruses 344 7.3.2 Infected host cells 347 7.4 The growth and control of viral populations 347 7.4.1 Virus productivity 347 7.4.2 Regulation of viral abundance 348 7.5 Control of host populations by aquatic viruses: impact on the microbial food web 351 7.5.1 Metabolic effects of viruses: reduction of algal primary productivity 351 7.5.2 Destruction of algal and bacterial populations 352 7.5.3 Viruses and the microbial loop 352 7.6 Cyanophages: viruses of blue-green algae 353 7.6.1 Classification and taxonomic characteristics 353 7.6.2 Infection of host cells 354 7.7 Phycoviruses: parasites of eukaryote algae 356 7.7.1 General characteristics 356 7.7.2 Host cell infection 358 7.7.3 Case Study: the infective life cycle of Chlorovirus 358 7.7.4 Ecological impact of phycoviruses 361 7.8 Virus infection of freshwater bacteria 362 7.8.1 General role of bacteriophages in the biology of freshwater bacteria 362 7.8.2 Bacteriophages in pelagic and benthic systems 362 7.8.3 Occurrence of free bacteriophages in aquatic systems 363 7.8.4 Incidence of bacterial infection 363 7.8.5 Temperate/virulent phage equilibrium and bacterial survival 365 7.8.6 Bacteriophage control of planktonic bacterial populations 367 7.8.7 Case study: viral lysis of bacteria in a eutrophic lake 367 7.8.8 Transduction: bacteriophage-mediated gene transfer between freshwater bacteria 369 7.8.9 Case study: transduction of plasmid and chromosomal DNA in Pseudomonas aeruginosa 370 8 Fungi and fungal-like organisms: aquatic biota with a mycelial growth form 373 A. ACTINOMYCETES, OOMYCETES, AND TRUE FUNGI 373 8.1 Fungi and fungal-like organisms: the mycelial growth habit 373 8.2 Actinomycetes 374 8.2.1 Taxonomic characteristics 374 8.2.2 Habitat 375 8.2.3 Nutrition 376 8.2.4 Competition with other microorganisms 376 8.3 Oomycetes 376 8.3.1 Oomycetes and true fungi 377 8.3.2 Taxonomic diversity 378 8.4 True fungi 379 8.4.1 Old and new terminology 379 8.4.2 Taxonomic diversity within the true fungi 380 B. FUNGI AS SAPROPHYTES AND PARASITES 383 8.5 Saprophytic activity of fungi 383 8.5.1 Colonization, growth, and fungal succession 384 8.5.2 Breakdown of leaf litter 385 8.5.3 Saprophytic fungi - chytrids and deuteromycetes 388 8.6 Parasitic activities of aquatic fungi 390 8.6.1 Parasitic and predatory deuteromycetes: fungi that attack small animals 390 8.6.2 Parasitic chytrids: highly specialized parasites of freshwater organisms 391 8.7 Fungal epidemics in the control of phytoplankton populations 394 8.7.1 Ecological significance 394 8.7.2 Case study: chytrid infection of the asterionella during an autumn diatom bloom 394 8.7.3 Net effect of infected and non-infected host cells 396 8.7.4 Factors affecting the development of fungal infection 397 9 Grazing activities in the freshwater environment: the role of protozoa and invertebrates 403 A. PROTOZOA 403 9.1 Introduction 403 9.1.1 Relative importance of protozoans, rotifers, and crustaceans in pelagic communities 403 9.1.2 Ecological role of protozoa 404 9.2 Protozoa, algae, and indeterminate groups 404 9.3 Taxonomic diversity of protozoa in the freshwater environment 405 9.3.1 Ciliates 405 9.3.2 Flagellate protozoa 409 9.3.3 Amoeboid protozoa 411 9.4 Ecological impact of protozoa: the pelagic environment 414 9.4.1 Positioning within the water column 414 9.4.2 Trophic interactions in the water column 415 9.5 Heterotrophic nanoflagellates: an integral component of planktonic communities 415 9.5.1 Enumeration of nanoflagellate populations in aquatic samples 416 9.5.2 Taxonomic composition of HNF communities 417 9.5.3 Abundance and control of flagellate populations 417 9.5.4 Nanoflagellate grazing rates and control of bacterial populations 418 9.5.5 Co-distribution of bacteria and protozoa within the water column 420 9.6 Ecological impact of protozoa: the benthic environment 421 9.6.1 Benthic microenvironments 421 9.6.2 Seasonal changes 423 9.6.3 Organic pollution 423 9.6.4 Sewage-treatment plants: activated sludge 424 B. GRAZING OF MICROBIAL POPULATIONS BY ZOOPLANKTON 425 9.7 General features of zooplankton: rotifers, cladocerans and copepods 425 9.7.1 Morphology and size 425 9.7.2 Reproduction and generation times 428 9.7.3 Predation of zooplankton 428 9.8 Grazing activity and prey selection 429 9.8.1 Seasonal succession in zooplankton feeding 429 9.8.2 Method of feeding 431 9.8.3 Selection of food by zooplankton 431 9.9 Grazing rates of zooplankton 434 9.9.1 Measurement 434 9.9.2 Factors affecting grazing rates 435 9.9.3 Diurnal variations in grazing activity 436 9.10 Effects of algal toxins on zooplankton 437 9.11 Biomass relationships between phytoplankton and zooplankton populations 439 9.11.1 Control of zooplankton populations 439 9.11.2 Biomass transfer 439 C. GRAZING OF BENTHIC MICROORGANISMS 439 9.12 Comparison of pelagic and benthic systems 440 9.13 Role of invertebrates in consuming river microorganisms 441 9.13.1 Grazing of periphyton biomass 442 9.13.2 Effects of grazing on periphyton community structure 443 10 Eutrophication: the microbial response to high nutrient levels 445 A. ORIGINS OF EUTROPHICATION 445 10.1 Nutrient status of freshwater environments: from oligotrophic to eutrophic systems 445 10.1.1 Eutrophic and oligotrophic lakes: definition of terms 446 10.1.2 Determinants of trophic status: location, morphology and hydrology 447 10.1.3 Artificial eutrophication: the impact of human activities 448 10.1.4 Eutrophication of rivers and streams 448 B. ECOLOGICAL EFFECTS OF EUTROPHICATION IN STANDING WATERS 450 10.2 General biological changes 450 10.2.1 The progression from oligotrophic to eutrophic waters 450 10.2.2 Effects of eutrophication on the water column of stratified lakes 453 10.2.3 Major changes in ecological balance: the breakdown of homeostasis 454 10.3 Biological assessment of water quality 455 10.3.1 Algal indicator groups 455 10.3.2 Case study: using the A/C (araphid pennate/centric) diatom ratio to assess eutrophication in Lake Tahoe (USA) 456 10.3.3 Indices of species diversity 457 10.4 Problems with intentional eutrophication: destabilization of fishpond ecosystems 457 10.4.1 Promotion of high productivity in fishponds 457 10.4.2 Destabilization and restoration of the ecosystem 458 C. THE GROWTH AND IMPACT OF ALGAL BLOOMS 459 10.5 Algal blooms and eutrophication 459 10.6 Formation of colonial blue-green algal blooms 461 10.6.1 General requirements for bloom formation 461 10.6.2 Competition with other algae 461 10.6.3 Case study: use of enclosure experiments to study factors affecting blue-green dominance 463 10.7 Environmental effects of blue-green blooms 463 10.7.1 General environmental changes 463 10.7.2 Specific effects on water quality 464 10.7.3 Production of toxins 464 D. CONTROL OF BLUE-GREEN ALGAE 466 10.8 Strategies for the control of blue-green algae 466 10.8.1 Nutrient limitation (bottom-up control) 467 10.8.2 Case study: restoration of water quality in Lake Washington, North West USA 468 10.8.3 Direct eradication 468 10.8.4 Top-down control of blue-green algae: the use of biomanipulation 469 10.9 Case study: top-down and bottom-up control of algal populations in the Broads Wetland Area (UK) 469 10.9.1 The eutrophication problem 469 10.9.2 Lake restoration 470 10.10 Biological control of blue-green algae 473 10.10.1 Biological control agents 473 10.10.2 Protocol for the development of biological control agents 475 10.10.3 Case study: potential protozoon control agents 477 10.10.4 Application of plant litter to control blue-green algae 478 10.11 Strategies for the control of blue-green algae in different water bodies 478 10.11.1 Integrated management policy 479 10.11.2 Case study: environmental monitoring at Hollingworth Lake, Greater Manchester (UK) 480 10.11.3 Specific remedial measures in different freshwater systems 482 References 485 Index 507
Library of Congress Subject Headings for this publication:
Freshwater microbiology.
Fresh Water -- microbiology.