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9 1.4.1 Summary of Foundation Concepts and Paradigms.. 36 3.11.1 Summary of Foundation Concepts and Paradigms.. 57 4.6.1 Summary of Foundation Concepts and Paradigms.. 72 5.11.1 Summary

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A Comprehensive Treatment

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A Comprehensive

Treatment

Michael C Newman William H Clements

CRC Press

Taylor & Francis Group Boca Raton London New York CRC Press is an imprint of the

Taylor Si Francis Group, an informa business

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CRC Press

Taylor & Francis Group

6000 Broken Sound Parkway NW, Suite 300

Boca Raton, FL 33487-2742

© 2008 by Taylor & Francis Group, LLC

CRC Press is an imprint of Taylor & Francis Group, an Informa business

No claim to original U.S Government works

Printed in the United States of America on acid-free paper

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International Standard Book Number-13:978-0-8493-3357-6 (Hardcover)

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Library of Congress Cataloging-in-Publication Data

Newman, Michael C

Ecotoxicology: a comprehensive treatment / Michael C Newman and William H Clements,

p.; cm

"A CRC title."

Includes bibliographical references and index

ISBN 978-0-8493-3357-6 (alk paper)

1 Toxicology—Environmental aspects I Clements, William H (William Henry), 1954- II Title

[DNLM: 1 Environmental Exposure—adverse effects 2 Environmental Pollutants—toxicity 3

Ecosystem 4 Population Dynamics WA 671 E196 2008]

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To Peg, Ben, and Ian (MCN)

To Diana for her endless support over the years (WHC)

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Do that which will render thee worthy of happiness

Critique of Pure Reason, I Kant 1781

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Preface xxv

Authors xxvii

I Hierarchical Ecotoxicology 1

Chapter 1 The Hierarchical Science of Ecotoxicology 3

1.1 An Overarching Context of Hierarchical Ecotoxicology 3

1.1.1 General 3

1.1.2 The Modified Janus Context 4

1.2 Reductionism versus Holism Debate 6

1.2.1 Reductionism versus Holism as a False Dichotomy 6

1.2.2 Microexplanation, Holism, and Macroexplanation 6

1.2.3 A Closer Look at Macroexplanation 7

1.3 Requirements in the Science of Ecotoxicology 8

1.3.1 General 8

1.3.2 Strongest Possible Inference 8

1.4 Summary 9

1.4.1 Summary of Foundation Concepts and Paradigms 9

References 10

II Organismal Ecotoxicology 11

Chapter 2 The Organismal Ecotoxicology Context 13

2.1 Overview 13

2.2 Organismal Ecotoxicology Defined 14

2.2.1 What Is Organismal Ecotoxicology? 14

2.3 The Value of Organismal Ecotoxicology Vantage 18

2.3.1 Tractability and Discreteness 18

2.3.2 Inferring Effects to or Exposure of Organisms with Suborganismal Metrics 18

2.3.3 Extrapolating among Individuals: Species, Size, Sex, and Other Key Qualities 19

2.3.4 Inferring Population Effects from Organismal Effects 19

2.3.5 Inferring Community Effects from Organismal Effects 20

2.3.6 Inferring Potential for Trophic Transfer from Bioaccumulation 21

2.4 Summary 21

References 21

Chapter 3 Biochemistry of Toxicants 23

3.1 Overview 23

3.2 DNA Modification 25

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3.3 Detoxification of Organic Compounds 25

3.3.1 Phase I Reactions 26

3.3.2 Phase II (Conjugative) Reactions 27

3.4 Metal Detoxification, Regulation, and Sequestration 28

3.5 Stress Proteins and Proteotoxicity 30

3.6 Oxidative Stress 31

3.7 Enzyme Dysfunction 32

3.8 Heme Biosynthesis Inhibition 32

3.9 Oxidative Phosphorylation Inhibition 35

3.10 Narcosis 35

3.11 Summary 36

3.11.1 Summary of Foundation Concepts and Paradigms 36

References 37

Chapter 4 Cells and Tissues 43

4.1 Overview 43

4.2 Cytotoxicity 43

4.2.1 Necrosis and Apoptosis 43

4.2.2 Types of Necrosis 44

4.2.3 Inflammation and Other Responses 47

4.3 Genotoxicity 50

4.3.1 Somatic and Genetic Risk 50

4.3.2 DNA Damage 52

4.3.3 Chromatids and Chromosomes 52

4.4 Cancer 53

4.4.1 Carcinogenesis 53

4.4.2 Cancer Latency 54

4.4.3 Threshold and Nonthreshold Models 55

4.5 Sequestration and Accumulation 55

4.5.1 Toxicants or Products of Toxicants 55

4.5.2 Cellular Materials as Evidence of Toxicant Damage 56

4.6 Summary 57

4.6.1 Summary of Foundation Concepts and Paradigms 57

References 57

Chapter 5 Organs and Organ Systems 63

5.1 Overview 63

5.2 General Integument 63

5.3 Organs Associated with Gas Exchange 65

5.3.1 Air Breathing 65

5.3.2 Water Breathing 66

5.4 Circulatory System 66

5.5 Digestive System 67

5.6 Liver and Analogous Organs of Invertebrates 68

5.7 Excretory Organs 69

5.8 Immune System 69

5.9 Endocrine System 70

5.10 Nervous, Sensory, and Motor-Related Organs and Systems 72

5.11 Summary 72

5.11.1 Summary of Foundation Concepts and Paradigms 72

References 73

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Chapter 6 Physiology 81

6.1 Overview 81

6.2 Ionic and Osmotic Regulation 82

6.3 Acid–Base Regulation 83

6.4 Respiration and General Metabolism 84

6.5 Bioenergetics 87

6.6 Plant-Related Processes 89

6.7 Summary 90

6.7.1 Summary of Foundation Concepts and Paradigms 90

References 91

Chapter 7 Bioaccumulation 95

7.1 Overview 95

7.2 Uptake 95

7.2.1 Cellular Mechanisms 95

7.2.2 Routes of Entry into Organisms 99

7.2.3 Factors Modifying Uptake 101

7.3 Biotransformation 104

7.4 Elimination 105

7.4.1 Hepatobiliary 106

7.4.2 Renal 106

7.4.3 Branchial 106

7.4.4 Other Elimination Mechanisms 107

7.5 Summary 107

7.5.1 Summary of Foundation Concepts and Paradigms 107

References 108

Chapter 8 Models of Bioaccumulation and Bioavailability 115

8.1 Overview 115

8.2 Bioaccumulation 115

8.2.1 Underlying Mechanisms 116

8.2.2 Assumptions of Models and Methods of Fitting Data 116

8.2.3 Rate Constant-Based Models 118

8.2.4 Clearance Volume-Based Models 122

8.2.5 Fugacity-Based Models 123

8.2.6 Physiologically Based Pharmacokinetic Models 125

8.2.7 Statistical Moments Formulations 125

8.3 Bioavailability 127

8.3.1 Conceptual Foundation: Concentration→Exposure→Realized Dose →Effect 127

8.3.2 Types and Estimation of Bioavailability 128

8.4 Summary 131

8.4.1 Summary of Foundation Concepts and Paradigms 131

References 132

Chapter 9 Lethal Effects 135

9.1 Overview 135

9.1.1 Distinct Dynamics Arising from Underlaying Mechanisms and Modes of Action 136

9.1.2 Lethality Differences among Individuals 140

9.1.2.1 Individual Effective Dose Hypothesis 141

9.1.2.2 Probabilistic Hypothesis 142

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9.1.3 Spontaneous and Threshold Responses 144

9.1.4 Hormesis 144

9.1.5 Toxicant Interactions 145

9.2 Quantifying Lethality 146

9.2.1 General 146

9.2.2 Dose or Concentration–Response Models Quantifying Lethality 146

9.2.3 Time–Response Models Quantifying Lethality 150

9.3 Lethality Prediction 154

9.3.1 Organic Compounds and the QSAR Approach 154

9.3.2 Metals and the QICAR Approach 156

9.4 Summary 157

9.4.1 Summary of Foundation Concepts and Paradigms 157

References 158

Chapter 10 Sublethal Effects 163

10.1 Overview 163

10.2 General Categories of Effects 166

10.2.1 Development and Growth 166

10.2.2 Reproduction 167

10.2.3 Behavior 167

10.2.4 Physiology 168

10.3 Quantifying Sublethal Effects 168

10.3.1 Hypothesis Testing and Point Estimation 169

10.3.1.1 Basic Concepts and Assumptions of Hypothesis Tests 175

10.3.1.2 Basic Concepts and Assumptions of Point Estimation Methods 179

10.4 Summary 179

10.4.1 Summary of Foundation Concepts and Paradigms 180

References 180

Chapter 11 Conclusion 189

11.1 General 189

11.2 Some Particularly Key Concepts 189

11.3 Concluding Remarks 191

III Population Ecotoxicology 193

Chapter 12 The Population Ecotoxicology Context 195

12.1 Population Ecotoxicology Defined 195

12.1.1 What Is a Population? 195

12.1.2 Definition of Population Ecotoxicology 196

12.2 The Need for Population Ecotoxicology 196

12.2.1 General 196

12.2.2 Scientific Merit 197

12.2.3 Practical Merit 199

12.3 Inferences within and between Biological Levels 203

12.3.1 Inferring Population Effects from Qualities of Individuals 204

12.3.2 Inferring Individual Effects from Qualities of Populations 204

12.3.3 Inferring Community Effects from Qualities of Populations 205

12.4 Summary 208

12.4.1 Summary of Foundation Concepts and Paradigms 208

References 208

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Chapter 13 Epidemiology: The Study of Disease in Populations 215

13.1 Foundation Concepts and Metrics in Epidemiology 215

13.1.1 Foundation Concepts 215

13.1.2 Foundation Metrics 218

13.1.3 Foundation Models Describing Disease in Populations 224

13.1.3.1 Accelerated Failure Time and Proportional Hazard Models 224

13.1.3.2 Binary Logistic Regression Model 227

13.2 Disease Association and Causation 228

13.2.1 Hill’s Nine Aspects of Disease Association 228

13.2.2 Strength of Evidence Hierarchy 232

13.3 Infectious Disease and Toxicant-Exposed Populations 235

13.4 Differences in Sensitivity within and among Populations 236

13.5 Summary 237

13.5.1 Summary of Foundation Concepts and Paradigms 237

References 238

Chapter 14 Toxicants and Simple Population Models 241

14.1 Toxicants Effects on Population Size and Dynamics 241

14.1.1 The Population-Based Paradigm for Ecological Risk 241

14.1.2 Evidence of the Need for the Population-Based Paradigm for Risk 242

14.2 Fundamentals of Population Dynamics 243

14.2.1 General 243

14.2.2 Projection Based on Phenomenological Models: Continuous Growth 244

14.2.3 Projection Based on Phenomenological Models: Discrete Growth 246

14.2.4 Sustainable Harvest and Time to Recovery 247

14.3 Population Stability 250

14.4 Spatial Distributions of Individuals in Populations 253

14.4.1 Describing Distributions: Clumped, Random, and Uniform 253

14.4.2 Metapopulations 254

14.4.2.1 Metapopulation Dynamics 254

14.4.2.2 Consequences to Exposed Populations 256

14.5 Summary 258

14.5.1 Summary of Foundation Concepts and Paradigms 258

References 259

Chapter 15 Toxicants and Population Demographics 263

15.1 Demography: Adding Individual Heterogeneity to Population Models 263

15.1.1 Structured Populations 263

15.1.2 Basic Life Tables 264

15.1.2.1 Survival Schedules 264

15.1.2.2 Mortality–Natality Tables 266

15.2 Matrix Forms of Demographic Models 270

15.2.1 Basics of Matrix Calculations 270

15.2.2 The Leslie Age-Structured Matrix Approach 272

15.2.3 The Lefkovitch Stage-Structured Matrix Approach 274

15.2.4 Stochastic Models 276

15.3 Summary 277

15.3.1 Summary of Foundation Concepts and Paradigms 277

References 278

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Chapter 16 Phenogenetics of Exposed Populations 281

16.1 Overview 281

16.1.1 The Phenotype Vantage 281

16.1.2 An Extreme Case Example 281

16.2 Toxicants and the Principle of Allocation (Concept of Strategy) 284

16.2.1 Phenotypic Plasticity and Norms of Reaction 286

16.2.2 Toxicants and Aging 289

16.2.2.1 Stress-Based Theories of Aging 290

16.2.2.2 Disposable Soma and Related Theories of Aging 290

16.2.3 Optimizing Fitness: Balancing Somatic Growth, Longevity, and Reproduction 291

16.3 Developmental Stability in Populations 294

16.4 Summary 297

16.4.1 Summary of Foundation Concepts and Paradigms 299

References 300

Chapter 17 Population Genetics: Damage and Stochastic Dynamics of the Germ Line 305

17.1 Overview 305

17.2 Direct Damage to the Germ Line 306

17.2.1 Genotoxicity 306

17.2.2 Repair of Genotoxic Damage 307

17.2.3 Mutation Rates and Accumulation 309

17.3 Indirect Change to the Germ Line 311

17.3.1 Stochastic Processes 311

17.3.2 Hardy–Weinberg Expectations 313

17.3.3 Genetic Drift 314

17.3.3.1 Effective Population Size 314

17.3.3.2 Genetic Bottlenecks 316

17.3.3.3 Balancing Drift and Mutation 317

17.3.4 Population Structure 317

17.3.4.1 The Wahlund Effect 317

17.3.4.2 Isolated and Semi-Isolated Subpopulations 320

17.3.5 Multiple Locus Heterozygosity and Individual Fitness 324

17.4 Genetic Diversity and Evolutionary Potential 326

17.5 Summary 326

17.5.1 Summary of Foundation Concepts and Paradigms 326

References 327

Chapter 18 Population Genetics: Natural Selection 331

18.1 Overview of Natural Selection 331

18.1.1 General 331

18.1.2 Viability Selection 334

18.1.3 Selection Components Associated with Reproduction 337

18.2 Estimating Differential Fitness and Natural Selection 340

18.2.1 Fitness, Relative Fitness, and Selection Coefficients 340

18.2.2 Heritability 343

18.3 Ecotoxicology’s Tradition of Tolerance 345

18.4 Summary 347

18.4.1 Summary of Foundation Concepts and Paradigms 347

References 348

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Chapter 19 Conclusion 353

19.1 Overview 353

19.2 Some Particularly Key Concepts 353

19.2.1 Epidemiology 353

19.2.2 Simple Models of Population Dynamics 354

19.2.3 Metapopulation Dynamics 354

19.2.4 The Demographic Approach 354

19.2.5 Phenogenetics Theory 355

19.2.6 Population Genetics: Stochastic Processes 355

19.2.7 Population Genetics: Natural Selection 356

19.3 Concluding Remarks 356

References 356

IV Community Ecotoxicology 359

Chapter 20 Introduction to Community Ecotoxicology 361

20.1 Definitions—Community Ecology and Ecotoxicology 361

20.1.1 Community Ecology 361

20.1.2 Community Ecotoxicology 362

20.2 Historical Perspective of Community Ecology and Ecotoxicology 362

20.2.1 Holism and Reductionism in Community Ecology and Ecotoxicology 363

20.2.2 Trophic Interactions in Community Ecology and Ecotoxicology 366

20.2.3 Importance of Experiments in Community Ecology and Ecotoxicology 366

20.3 Are Communities More Than the Sum of Individual Populations? 367

20.3.1 The Need to Understand Indirect Effects of Contaminants 367

20.4 Communities within the Hierarchy of Biological Organization 370

20.5 Contemporary Topics in Community Ecotoxicology 372

20.5.1 The Need for an Improved Understanding of Basic Community Ecology 372

20.5.2 Development and Application of Improved Biomonitoring Techniques 372

20.5.3 Application of Contemporary Food Web Theory to Ecotoxicology 373

20.5.4 The Need for Improved Experimental Approaches 374

20.5.5 Influence of Global Atmospheric Stressors on Community Responses to Contaminants 374

20.6 Summary 375

20.6.1 Summary of Foundation Concepts and Paradigms 375

References 376

Chapter 21 Biotic and Abiotic Factors That Regulate Communities 379

21.1 Characterizing Community Structure and Organization 379

21.1.1 Colonization and Community Structure 381

21.1.2 Definitions of Species Diversity 381

21.2 Changes in Species Diversity and Composition along Environmental Gradients 382

21.2.1 Global Patterns of Species Diversity 383

21.2.2 Species–Area Relationships 385

21.2.3 Assumptions about Equilibrium Communities 387

21.3 The Role of Keystone Species in Community Regulation 388

21.3.1 Identifying Keystone Species 389

21.4 The Role of Species Interactions in Community Ecology and Ecotoxicology 391

21.4.1 Definitions 391

21.4.2 Experimental Designs for Studying Species Interactions 392

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21.4.3 The Influence of Contaminants on Predator–Prey Interactions 393

21.4.4 The Influence of Contaminants on Competitive Interactions 397

21.5 Environmental Factors and Species Interactions 399

21.5.1 Environmental Stress Gradients 400

21.6 Summary 401

21.6.1 Summary of Foundation Concepts and Paradigms 402

References 403

Chapter 22 Biomonitoring and the Responses of Communities to Contaminants 409

22.1 Biomonitoring and Biological Integrity 409

22.2 Conventional Approaches 410

22.2.1 Indicator Species Concept 410

22.3 Biomonitoring and Community-Level Assessments 411

22.3.1 Species Abundance Models 411

22.3.2 The Use of Species Richness and Diversity to Characterize Communities 415

22.3.2.1 Species Richness 415

22.3.2.2 Species Diversity 417

22.3.2.3 Species Evenness 418

22.3.2.4 Limitations of Species Richness and Diversity Measures 418

22.3.3 Biotic Indices 420

22.4 Development and Application of Rapid Bioassessment Protocols 423

22.4.1 Application of Qualitative Sampling Techniques 425

22.4.2 Subsampling and Fixed-Count Sample Processing 425

22.4.3 Pooling Samples 426

22.4.4 Relaxed Taxonomic Resolution 427

22.4.5 The Application of Species Traits in Biomonitoring 429

22.5 Regional Reference Conditions 430

22.6 Integrated Assessments of Biological Integrity 431

22.7 Limitations of Biomonitoring 432

22.7.1 Summary 434

22.7.1.1 Summary of Foundation Concepts and Paradigms 434

References 435

Chapter 23 Experimental Approaches in Community Ecology and Ecotoxicology 439

23.1 Experimental Approaches in Basic Community Ecology 439

23.1.1 The Transition from Descriptive to Experimental Ecology 439

23.1.2 Manipulative Experiments in Rocky Intertidal Communities 442

23.1.3 Manipulative Studies in More Complex Communities 442

23.1.4 Types of Experiments in Basic Community Ecology 443

23.2 Experimental Approaches in Community Ecotoxicology 444

23.3 Microcosms and Mesocosms 445

23.3.1 Background and Definitions 445

23.3.2 Design Considerations in Microcosm and Mesocosm Studies 447

23.3.2.1 Source of Organisms in Microcosm Experiments 447

23.3.2.2 Spatiotemporal Scale of Microcosm and Mesocosm Experiments 448

23.3.2.3 The Influence of Seasonal Variation on Community Responses 450

23.3.3 Statistical Analyses of Microcosm and Mesocosm Experiments 450

23.3.4 General Applications of Microcosms and Mesocosms 451

23.3.4.1 The Use of Mesocosms for Pesticide Registration 452

23.3.4.2 Development of Concentration–Response Relationships 452

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23.3.4.3 Investigation of Stressor Interactions 453

23.3.4.4 Influence of Environmental and Ecological Factors on Community Responses 454

23.3.4.5 Species Interactions 455

23.3.4.6 Applications in Terrestrial Systems 455

23.3.5 Summary 457

23.4 Whole Ecosystem Manipulations 457

23.4.1 Examples of Ecosystem Manipulations: Aquatic Communities 458

23.4.1.1 Experimental Lakes Area (ELA) 458

23.4.1.2 Coweeta Hydrologic Laboratory 459

23.4.1.3 Summary 459

23.4.2 Examples of Ecosystem Manipulations: Avian and Mammalian Communities 460

23.4.3 Limitations of Whole Ecosystem Experiments 462

23.5 What Is the Appropriate Experimental Approach for Community Ecotoxicology? 464

23.5.1 Questions of Spatiotemporal Scale 464

23.5.2 Integrating Descriptive and Experimental Approaches 464

23.6 Summary 465

23.6.1 Summary of Foundation Concepts and Paradigms 466

References 467

Chapter 24 Application of Multimetric and Multivariate Approaches in Community Ecotoxicology 473

24.1 Introduction 473

24.1.1 Comparison of Multimetric and Multivariate Approaches 474

24.2 Multimetric Indices 475

24.2.1 Multimetric Approaches for Terrestrial Communities 477

24.2.2 Limitations of Multimetric Approaches 478

24.3 Multivariate Approaches 479

24.3.1 Similarity Indices 479

24.3.2 Ordination 481

24.3.3 Discriminant and Cluster Analysis 486

24.3.4 Application of Multivariate Methods to Laboratory Data 488

24.3.5 Taxonomic Aggregation in Multivariate Analyses 490

24.4 Summary 491

24.4.1 Summary of Foundation Concepts and Paradigms 491

References 492

Chapter 25 Disturbance Ecology and the Responses of Communities to Contaminants 497

25.1 The Importance of Disturbance in Structuring Communities 497

25.1.1 Disturbance and Equilibrium Communities 498

25.1.2 Resistance and Resilience Stability 499

25.1.3 Pulse and Press Disturbances 500

25.2 Community Stability and Species Diversity 502

25.3 Relationship between Natural and Anthropogenic Disturbance 504

25.3.1 The Ecosystem Distress Syndrome 505

25.3.2 The Intermediate Disturbance Hypothesis 506

25.3.3 Subsidy–Stress Gradients 508

25.4 Contemporary Hypotheses to Explain Community Responses to Anthropogenic Disturbance 509

25.4.1 Pollution-Induced Community Tolerance 510

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25.5 Biotic and Abiotic Factors That Influence Community Recovery 512

25.5.1 Cross-Community Comparisons of Recovery 514

25.5.2 Importance of Long-Term Studies for Documenting Recovery 515

25.5.3 Community-Level Indicators of Recovery 515

25.5.4 Community Characteristics that Influence Rate of Recovery 519

25.6 Influence of Environmental Variability on Resistance and Resilience 521

25.7 Quantifying the Effects of Compound Perturbations 523

25.7.1 Sensitivity of Communities to Novel Stressors 523

25.8 Summary 526

25.8.1 Summary of Foundation Concepts and Paradigms 526

References 528

Chapter 26 Community Responses to Global and Atmospheric Stressors 533

26.1 Introduction 533

26.2 CO2and Climate Change 534

26.2.1 Facts and Evidence 535

26.2.2 Carbon Cycles and Sinks 537

26.2.3 The Mismatch between Climate Models and Ecological Studies 539

26.2.4 Paleoecological Studies of CO2and Climate Change 540

26.2.5 Effects of Climate Change on Terrestrial Vegetation 541

26.2.6 Ecological Responses to CO2Enrichment 543

26.2.7 Effects of Climate Change on Terrestrial Animal Communities 544

26.2.8 Effects of Climate Change on Freshwater Communities 546

26.2.9 Effects of Climate Change on Marine Communities 549

26.2.10 Conclusions 551

26.3 Stratospheric Ozone Depletion 552

26.3.1 Methodological Approaches for Manipulating UVR 554

26.3.2 The Effects of UVR on Marine and Freshwater Plankton 554

26.3.2.1 Direct and Indirect Effects of UV-B Radiation 555

26.3.3 Responses of Benthic Communities 556

26.3.4 Responses of Terrestrial Plant Communities 557

26.3.5 Biotic and Abiotic Factors That Influence UV-B Effects on Communities 558

26.3.5.1 Dissolved Organic Materials 558

26.3.5.2 Location 559

26.3.5.3 Interspecific and Intraspecific Differences in UV-B Tolerance 560

26.3.5.4 Interactions with Other Stressors 561

26.4 Acid Deposition 562

26.4.1 Descriptive Studies of Acid Deposition Effects in Aquatic Communities 562

26.4.2 Episodic Acidification 564

26.4.3 Experimental Studies of Acid Deposition Effects in Aquatic Communities 565

26.4.4 Recovery of Aquatic Ecosystems from Acidification 566

26.4.5 Effects of Acid Deposition on Forest Communities 567

26.4.6 Indirect Effects of Acidification on Terrestrial Wildlife 569

26.5 Interactions among Global Atmospheric Stressors 569

26.6 Summary 571

26.6.1 Summary of Foundation Concepts and Paradigms 572

References 574

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Chapter 27 Effects of Contaminants on Trophic Structure and Food Webs 581

27.1 Introduction 581

27.2 Basic Principles of Food Web Ecology 582

27.2.1 Historical Perspective of Food Web Ecology 582

27.2.2 Descriptive, Interactive, and Energetic Food Webs 583

27.2.3 Contemporary Questions in Food Web Ecology 584

27.2.4 Trophic Cascades 587

27.2.5 Limitations of Food Web Studies 590

27.2.6 Use of Radioactive and Stable Isotopes to Characterize Food Webs 592

27.3 Effects of Contaminants on Food Chains and Food Web Structure 592

27.3.1 Interspecific Differences in Contaminant Sensitivity 593

27.3.2 Indirect Effects of Contaminant Exposure on Feeding Habits 594

27.3.3 Alterations in Energy Flow and Trophic Structure 595

27.4 Summary 597

27.4.1 Summary of Foundation Concepts and Paradigms 597

References 598

Chapter 28 Conclusions 603

28.1 General 603

28.2 Some Particularly Key Concepts 603

28.2.1 Improvements in Experimental Techniques 603

28.2.2 Use of Multimetric and Multivariate Approaches to Assess Community-Level Responses 604

28.2.3 Disturbance Ecology and Community Ecotoxicology 604

28.2.4 An Improved Understanding of Trophic Interactions 605

28.2.5 Interactions between Contaminants and Global Atmospheric Stressors 606

28.3 Summary 607

28.3.1 Summary of Foundation Concepts and Paradigms 607

References 608

V Ecosystem Ecotoxicology 611

Chapter 29 Introduction to Ecosystem Ecology and Ecotoxicology 613

29.1 Background and Definitions 613

29.1.1 The Spatial Boundaries of Ecosystems 614

29.1.2 Contrast of Energy Flow and Materials Cycling 614

29.1.3 Community Structure, Ecosystem Function and Stability 615

29.2 Ecosystem Ecology and Ecotoxicology: A Historical Context 615

29.2.1 Early Development of the Ecosystem Concept 616

29.2.2 Quantification of Energy Flow through Ecosystems 617

29.2.3 The International Biological Program and the Maturation of Ecosystem Science 618

29.3 Challenges to the Study of Whole Systems 619

29.3.1 Temporal Scale 619

29.4 The Role of Ecosystem Theory 621

29.4.1 Succession Theory and the Strategy of Ecosystem Development 621

29.4.2 Hierarchy Theory and the Holistic Perspective of Ecosystems 622

29.5 Recent Developments in Ecosystem Science 623

29.5.1 General Methodological Approaches 624

29.5.2 The Importance of Multidisciplinary Research in Ecosystem Ecology and Ecotoxicology 625

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