Biogeochemistry of Wetlands Science and Applicatio

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Biogeochemistry of Wetlands: Science and Applications
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DOI: 10.1201/9780429155833
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Biogeochemistry of Wetlands
The globally important nature of wetland ecosystems has led to their increased protection and
restoration as well as their use in engineered systems. Underpinning the beneficial functions of
wetlands are a unique suite of physical, chemical, and biological processes that regulate elemental
cycling in soils and the water column. This book provides an in-depth coverage of these wetland
biogeochemical processes related to the cycling of macroelements, including carbon, nitrogen,
phosphorus, and sulfur; secondary and trace elements; and toxic organic compounds.
In this synthesis, the authors combine more than 100 years of experience studying wetlands and
biogeochemistry to look inside the black box of elemental transformations in wetland ecosystems.
This new edition is updated throughout to include more topics and provide an integrated view of
the coupled nature of biogeochemical cycles in wetland systems. The influence of the elemental
cycles is discussed at a range of scales in the context of environmental change including climate,
sea level rise, and water quality. Frequent examples of key methods and major case studies are also
included to help the reader extend the basic theories for application in their own system. Some of
the major topics discussed are:
• Flooded soil and sediment characteristics

• Aerobic-anaerobic interfaces
• Redox chemistry in flooded soil and sediment systems
• Anaerobic microbial metabolism
• Plant adaptations to reducing conditions
• Regulators of organic matter decomposition and accretion
• Major nutrient sources and sinks
• Greenhouse gas production and emission
• Elemental flux processes
• Remediation of contaminated soils and sediments
• Coupled C-N-P-S processes
• Consequences of environmental change in wetlands
The book provides the foundation for a basic understanding of key biogeochemical processes and its
applications to solve real-world problems. It is detailed, but also assists the reader with box inserts,
artfully designed diagrams, and summary tables all supported by numerous current references. This
book is an excellent resource for senior undergraduates and graduate students studying ecosystem
biogeochemistry with a focus in wetlands and aquatic systems.
Biogeochemistry of Wetlands
Science and Applications
Second Edition
K. Ramesh Reddy, Ronald D. DeLaune and

Patrick W. Inglett
Cover image: Shutterstock
Second edition published 2023
by CRC Press
6000 Broken Sound Parkway NW, Suite 300, Boca Raton, FL 33487–2742
and by CRC Press
4 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN
CRC Press is an imprint of Taylor & Francis Group, LLC
© 2023 K. Ramesh Reddy, Ronald D. DeLaune and Patrick W. Inglett
First edition published by CRC Press 2008
Reasonable efforts have been made to publish reliable data and information, but the
author and publisher cannot assume responsibility for the validity of all materials or the
consequences of their use. The authors and publishers have attempted to trace the copyright
holders of all material reproduced in this publication and apologize to copyright holders if
permission to publish in this form has not been obtained. If any copyright material has not
been acknowledged please write and let us know so we may rectify in any future reprint.
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means, now known or hereafter invented, including photocopying, microfilming, and
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Trademark notice: Product or corporate names may be trademarks or registered trademarks
and are used only for identification and explanation without intent to infringe.
ISBN: 978-1-4987-6455-1 (hbk)
ISBN: 978-1-032-32235-3 (pbk)
ISBN: 978-0-429-15583-3 (ebk)
DOI: 10.1201/9780429155833
Typeset in Times New Roman
by Apex CoVantage, LLC
Contents Contents Biogeochemistry
Preface��xv
Acknowledgments��xvii
Authors��xix
Chapter 1 Introduction��1
Chapter 2 Basic Concepts and Terminology��9

2.1 Introduction��9
2.2 Chemistry��9
2.2.1 Aqueous Chemistry�� 9
2.2.1.1 Concentration Units�� 10
2.2.2 Acids and Bases�� 10
2.2.3 Equilibrium Constant�� 11
2.2.4 Thermodynamics�� 11
2.2.4.1 Influence of pH�� 14
2.2.5 Oxidation–Reduction Reactions�� 15
2.2.5.1 Oxidation–Reduction�� 15
2.2.5.2 Oxidation State or Number�� 15
2.2.6 Balancing Oxidation–Reduction Reactions�� 16
2.3 Biology�� 18
2.3.1 Microbial Cell�� 18
2.3.2 Microbial Classification�� 18
2.3.3 Chemistry of Biological Molecules�� 19
2.3.4 Metabolic Reactions�� 19
2.3.5 Enzymes�� 20
2.3.6 Biochemical Kinetics�� 20
2.4 Isotopes�� 21
2.4.1 Radioactive Isotopes and Decay�� 21
2.4.2 Half-Life�� 21
2.4.3 Stable Isotopes�� 22
2.5 Terminology in Soil Science�� 24
2.5.1 Master Soil Horizon�� 24
2.5.2 Properties Used in Soil Description�� 25
2.5.3 Soil Taxonomy�� 25
2.5.4 Physical Properties�� 28
2.5.5 Chemical Properties�� 28
2.6 Units�� 29
Study Questions�� 30
Further Readings��31
Chapter 3 Biogeochemical Characteristics�� 33

3.1 Introduction�� 33
3.2 Types of Wetlands�� 38
3.2.1 Coastal Wetlands�� 38
3.2.2 Inland Wetlands�� 39
v
vi Contents
3.3 Wetland Hydrology�� 41

3.4 Wetland Soils�� 42
3.4.1 Physical Characteristics�� 43
3.4.2 Biochemical Characteristics�� 45
3.4.3 Biological Characteristics�� 48
3.5 Wetland Vegetation�� 48
3.6 Biogeochemical Features of Wetlands�� 49
3.6.1 Presence of Molecular Oxygen in Restricted Zones�� 49
3.6.2 Sequential Reduction of Other Inorganic Electron Acceptors�� 50
3.6.3 Aerobic Soil–Floodwater Interface�� 52
3.6.4 Exchanges at the Soil–Water Interface�� 52
3.6.5 Presence of Hydrophytic Vegetation�� 54
3.7 Types of Wetland/Hydric Soils�� 55
3.7.1 Waterlogged Mineral Soils�� 55
3.7.2 Organic Soils (Histosols)�� 59
3.7.3 Marsh Soils�� 61
3.7.4 Paddy Soils�� 62
3.7.5 Subaqueous Soils�� 62
3.7.6 Hydric Soils�� 63
3.8 Summary��64
Chapter 4 Electrochemical Properties�� 69

4.2 Theoretical Relationships�� 72
4.2.1 E° vs. log K��80
4.2.2 pe vs. Eh�� 81
4.3 Measurement of Eh��84
4.4 Eh–pH Relationships�� 85
4.5 Buffering of Redox Potential (Poise)�� 89
4.6 Measurement of Redox Potentials�� 89
4.6.1 Construction of Platinum Electrodes�� 89
4.6.2 Standardization of Electrodes��92
4.6.3 Redox Potentials in Soils�� 93
4.7 pH�� 97
4.7.1 Soil pH�� 98
4.7.2 Floodwater pH�� 100
4.7.3 pH Effects�� 101
4.8 Redox Couples in Wetlands�� 103
4.8.1 Intensity�� 103
4.8.2 Capacity�� 104
4.9 Redox Gradients in Soils�� 105
4.10 Specific Conductance�� 112
4.11 Summary�� 112
Further Readings�� 116
Contents vii
Chapter 5 Carbon��117
5.1 Introduction��117
5.2 Major Components of the Carbon Cycle in Wetlands�� 119
5.2.1 Plant Biomass Carbon (Net Primary Productivity)�� 120
5.2.2 Particulate Organic Matter (Detrital and Soil)�� 122
5.2.3 Microbial Biomass Carbon�� 123
5.2.4 Dissolved Organic Matter�� 124
5.2.5 Gaseous Forms of Carbon�� 125
5.3 Organic Matter Accumulation�� 126
5.4 Characteristics of Detritus and Soil Organic Matter�� 128
5.4.1 Non-Humic Substances�� 130
5.4.1.1 Carbohydrates�� 130
5.4.2 Phenolic Substances�� 131
5.4.3 Humic Substances�� 134
5.5 Decomposition�� 138
5.5.1 Leaching and Fragmentation�� 138
5.5.2 Photolysis�� 139
5.5.3 Extracellular Enzyme Hydrolysis�� 141
5.5.4 Catabolic Activity�� 146
5.5.4.1 Aerobic Catabolism�� 148
5.5.4.2 Anaerobic Catabolism�� 151
5.5.4.3 Aerobic vs. Anaerobic Catabolism�� 160
5.6 Organic Matter Turnover�� 162
5.6.1 Decomposition Rates�� 162
5.7 Regulators of Organic Matter Decomposition�� 166
5.7.1 Quality and Quantity of Organic Matter�� 167
5.7.2 Microbial Communities and Biomass�� 170
5.7.3 Water Table or Soil Aeration Status�� 171
5.7.4 Availability of Electron Acceptors with Higher
Reduction Potentials�� 174
5.7.5 Nutrient Availability�� 177
5.7.6 Temperature�� 180
5.8 Environmental and Ecological Significance�� 186
5.9 Functions of Organic Matter in Soils�� 193
5.10 Summary�� 197
Chapter 6 Oxygen�� 203

6.2 Soil Gases��204
6.3 Oxygen–H2O Redox Couple��209
6.3.1 Oxygen Diffusion Rate�� 213
6.3.2 Soil Oxygen Content�� 216
6.4 Sources of Oxygen�� 222
6.5 Aerobic-A naerobic Interfaces�� 225
6.6 Oxygen Consumption�� 228
6.6.1 Oxygen as a Reactant�� 228
6.6.2 Oxygen as an Electron Acceptor�� 229
viii Contents
6.7 Summary�� 235

Chapter 7 Adaptation of Plants to Soil Anaerobiosis�� 239

7.2 Distribution of Wetland Plants�� 241
7.3 Mechanisms of Flood Tolerance�� 242
7.3.1 Metabolic Adaptations�� 242
7.3.2 Morphological/Anatomical Adaptations�� 245
7.3.2.1 Roots�� 245
7.3.2.2 Pneumatophores�� 245
7.3.2.3 Lenticels�� 245
7.3.2.4 Intercellular Airspaces�� 245
7.3.3 Aerenchyma Formation��246
7.3.4 Intercellular Oxygen Concentration�� 249
7.4 Mechanisms of Oxygen Movement in Wetland Plants�� 251
7.4.1 Diffusion�� 251
7.4.2 Mass Flow�� 252
7.5 Oxygen Release by Plants�� 260
7.6 Measurement of Radial Oxygen Loss�� 261
7.7 Soil Phytotoxic Accumulation Effects on Plant Growth��264
7.7.1 Greenhouse Gas Emissions: Methane�� 267
7.7.2 Greenhouse Gas Emissions: Nitrous Oxide�� 269
7.8 Oxidizing Power of Plant Roots�� 270
7.8.1 Root Iron Plaque Formation�� 271
7.9 Effect of Intensity and Capacity of Soil Reduction on Wetland
Plant Functions�� 272
7.9.1 Effect of Soil Reduction Intensity�� 273
7.9.2 Relationship of Reduction Intensity with Root Porosity
and Radial Oxygen Loss�� 275
7.9.3 Effect of Soil Reduction Intensity on Nutrient Uptake�� 276
7.9.4 Soil Reduction Capacity Effects on Carbon Assimilation
and Radial Oxygen Loss�� 276
7.10 Summary�� 278
Chapter 8 Nitrogen�� 281

8.2 Forms of Nitrogen�� 281
8.2.1 Inorganic Nitrogen�� 281
8.2.2 Organic Nitrogen�� 282
8.3 Major Storage Compartments�� 282
8.3.1 Plant Biomass Nitrogen��284
8.3.2 Particulate Organic Nitrogen��284
8.3.3 Microbial Biomass Nitrogen�� 285
8.3.4 Dissolved Organic Nitrogen�� 285
Contents ix
8.3.5 Inorganic Forms of Nitrogen�� 285

8.3.6 Gaseous Forms of Nitrogen�� 286
8.4 Redox Transformations of Nitrogen�� 286
8.5 Nitrogen Fixation�� 287
8.5.1 Regulators of Dinitrogen Fixation�� 289
8.5.2 Nitrogen Fixation Rates�� 292
8.6 Nitrogen Assimilation by Vegetation�� 293
8.7 Organic Nitrogen Accumulation�� 296
8.8 Mineralization of Organic Nitrogen�� 297
8.8.1 Chemical Composition of Organic Nitrogen�� 298
8.8.2 C:N Ratio Concept�� 305
8.8.3 Microbial Degradation of Organic Nitrogen�� 308
8.8.4 Regulators of Organic Nitrogen Mineralization�� 312
8.9 Ammonia Adsorption–Desorption�� 314
8.10 Ammonia Fixation�� 316
8.11 Ammonia Volatilization��317
8.11.1 Physicochemical Reaction�� 318
8.11.2 Regulators of Ammonia Volatilization�� 320
8.12 Aerobic Ammonia Oxidation�� 323
8.12.1 Chemoautotrophic Prokaryotes�� 324
8.12.2 Methane-Oxidizing Bacteria�� 325
8.12.3 Heterotrophic Bacteria and Fungi�� 326
8.12.4 Regulators of Ammonium Oxidation�� 326
8.13 Anaerobic Ammonium Oxidation�� 328
8.13.1 Other Processes of Anaerobic Ammonium Oxidation�� 330
8.14 Nitrate Reduction�� 332
8.14.1 Denitrification�� 334
8.14.2 Nitrifier Denitrification�� 337
8.14.3 Aerobic Denitrification�� 338
8.14.4 Chemodenitrification�� 339
8.14.5 Dissimilatory Nitrate Reduction to Ammonia��340
8.14.6 Regulators of Nitrate Reduction�� 341
8.14.7 Nitrate Reduction Rates in Wetlands and Aquatic Systems��344
8.15 Nitrogen Processing by Wetlands��346
8.15.1 Ammonium Flux�� 347
8.15.2 Nitrate Flux�� 347
8.17 Summary�� 352
Chapter 9 Phosphorus�� 355

9.2 Phosphorus Accumulation in Soils�� 357
9.2.1 Why Does Phosphorus Added to Wetlands Accumulate
in Soils?�� 357
9.3 Phosphorus Forms in the Water Column and Soil�� 359
9.3.1 Phosphorus Speciation�� 359
9.3.2 Water Column�� 360
x Contents
9.3.2 Soil�� 362

9.4 Inorganic Phosphorus��364
9.5 Phosphorus Sorption by Soils�� 370
9.5.1 Adsorption–Desorption�� 373
9.5.1.1 Isotherm Concepts�� 375
9.5.2 Phosphorus Sorption Isotherms�� 377
9.5.2.1 Linear Equation�� 378
9.5.2.2 Freundlich Equation�� 378
9.5.2.3 Langmuir Equation�� 379
9.5.2.4 Single-Point Isotherm�� 379
9.5.2.5 Quantity (Q)/Intensity (I) Relationships�� 380
9.5.3 Precipitation and Dissolution�� 380
9.5.4 Regulators of Phosphorus Retention and Release�� 383
9.6 Organic Phosphorus�� 387
9.6.1 Forms of Organic Phosphorus�� 387
9.6.2 Chemical Characterization of Organic Phosphorus�� 391
9.7 Phosphorus Uptake and Storage in Biotic Communities�� 394
9.7.1 Microorganisms�� 394
9.7.2 Periphyton�� 396
9.7.3 Vegetation�� 396
9.8 Mineralization of Organic Phosphorus��400
9.8.1 Abiotic Degradation and Stabilization of Organic Phosphorus�� 401
9.8.1.1 Leaching of Soluble Organic Phosphorus�� 401
9.8.1.2 Noncatalyzed Hydrolysis of Phosphate Esters�� 401
9.8.1.3 Photolysis��402
9.8.1.4 Stabilization of Organic Phosphorus��402
9.8.2 Enzymatic Hydrolysis of Organic Phosphorus��402
9.8.2.1 Phosphatases or Monoesterases��404
9.8.2.2 Phosphodiesterases�� 405
9.8.3 Microbial Activities and Phosphorus Release�� 405
9.8.4 Regulators of Organic Phosphorus Mineralization�� 410
9.9 Biotic and Abiotic Interactions on Phosphorus Mobilization�� 412
9.9.1 Phosphorus–Iron–Sulfur Interactions�� 412
9.9.2 Periphyton–Phosphate Interactions�� 414
9.9.3 Biotic and Abiotic Interactions of Fe and Ca with Phosphorus�� 417
9.9.4 Gaseous Loss of Phosphorus�� 418
9.10 Phosphorus Exchange between Soil and Overlying Water Column�� 419
9.11 Phosphorus Memory by Soils and Sediments�� 421
9.12 Summary�� 425
Chapter 10 Iron and Manganese�� 429

10.2 Storage and Distribution�� 429
10.3 Eh–pH Relationships�� 432
10.3.1 Iron�� 433
10.3.2 Manganese�� 435
Contents xi
10.4 Reduction of Iron and Manganese�� 435

10.4.1 Microbial Communities�� 437
10.4.2 Biotic and Abiotic Reduction�� 439
10.4.2.1 Biotic Reduction��440
10.4.2.2 Abiotic Reduction�� 441
10.4.3 Forms of Iron and Manganese��446
10.4.3.1 Iron��446
10.4.3.2 Manganese��448
10.4.3.3 Complexation of Iron and Manganese with
Dissolved Organic Matter�� 449
10.4.3.4 Mobile and Immobile Pools of Iron and Manganese�� 449
10.5 Oxidation of Iron and Manganese�� 452
10.5.2 Biotic and Abiotic Oxidation�� 454
10.5.2.1 Iron�� 454
10.5.2.2 Manganese�� 458
10.6 Mobility of Iron and Manganese�� 458
10.7 Ecological Significance�� 462
10.7.1 Nutrient Regeneration/Immobilization�� 462
10.7.1.1 Organic Matter Decomposition and Nutrient Release�� 462
10.7.1.2 Phosphorus Release or Retention�� 463
10.7.1.3 Coprecipitation of Trace Elements with Iron and
Manganese Oxides��464
10.7.1.4 Siderophores and Complexation of Iron and
Manganese Oxides��464
10.7.2 Ferromanganese Nodules�� 465
10.7.3 Root Plaque Formation��466
10.7.4 Wetting and Drying: Hydrologic Fluctuations��466
10.7.5 Ferrolysis�� 467
10.7.6 Methane Emissions�� 467
10.8 Summary�� 469
Chapter 11 Sulfur�� 473

11.2 Major Storage Compartments�� 474
11.3 Forms of Sulfur�� 474
11.4 Oxidation–Reduction of Sulfur�� 477
11.5 Assimilatory Sulfate and Elemental Sulfur Reduction�� 480
11.6 Mineralization of Organic Sulfur�� 482
11.7 Electron Acceptor–Reduction of Inorganic Sulfur�� 483
11.7.1 Dissimilatory Sulfate Reduction��484
11.7.2 Role of Sulfur in Energy Flow�� 488
11.7.3 Measurement of Sulfate Reduction in Wetland Soils�� 488
11.7.4 Regulators of Sulfate Reductions�� 490
11.8 Sulfide Toxicity�� 492
11.9 Electron Donor–Oxidation of Sulfur Compounds�� 493
xii Contents
11.10 Biogenic Emission of Reduced Sulfur Gases�� 496

11.11 Sulfur–Metal Interactions�� 498
11.12 Exchange between Soil and Water Column�� 503
11.13 Sulfur Sinks��504
11.15 Summary�� 506
Chapter 12 Metals/Metalloids�� 509

12.2 Biogeochemical Regulators of Metal Availability and
Transformation�� 509
12.2.1 Sorption and Precipitation�� 511
12.2.2 Interaction with Organic Matter�� 511
12.2.3 Interaction with Clay Minerals�� 512
12.2.4 Biotic Transformations�� 513
12.2.5 Redox Potential and pH of Soils and Sediments�� 513
12.3 Mercury–Methyl Mercury�� 515
12.4 Arsenic�� 519
12.5 Copper�� 525
12.6 Zinc�� 528
12.7 Selenium�� 529
12.8 Chromium�� 532
12.9 Cadmium�� 535
12.10 Lead�� 537
12.11 Nickel�� 538
12.12 Summary��540
Chapter 13 Toxic Organic Compounds�� 543

13.2 Abiotic Pathways�� 554
13.2.1 Redox Potential–pH�� 554
13.2.2 Hydrolysis�� 555
13.2.3 Sorption to Suspended Solids and the Substrate Bed�� 555
13.2.3.1 Effect of Colloidal Organic Matter in Surface
Water on Sorption in Wetlands�� 557
13.2.4 Photolysis�� 558
13.3 Biotic Pathways�� 559
13.3.1 Acclimation�� 560
13.3.2 Biodegradation�� 560
13.3.3 Cometabolism�� 560
13.3.4 Microbial Accumulation�� 560
13.3.5 Polymerization and Conjugation�� 561
13.4 Metabolism of Organic Compounds�� 561
13.4.1 Hydrolysis�� 561
13.4.2 Oxidation�� 562
Contents xiii
13.4.2.1 Hydroxylation��564
13.4.2.2 Dealkylation��564
13.4.2.3 β-Oxidation��564
13.4.2.4 Decarboxylation��564
13.4.2.5 Cleavage of Ether Linkage�� 565
13.4.2.6 Epoxidation�� 565
13.4.2.7 Oxidative Coupling�� 565
13.4.2.8 Aromatic Ring Cleavage�� 565
13.4.2.9 Heterocyclic Ring Cleavage�� 565
13.4.2.10 Sulfoxidation�� 565
13.5.3 Reduction�� 566
13.4.3.1 Reductive Dehalogenation�� 566
13.4.4 Synthesis�� 566
13.5 Plant and Microbial Uptake�� 568
13.6 Transport Processes�� 568
13.6.1 Exchange between Soil and Water Column�� 568
13.6.2 Settling and Burial of Particulate Contaminants�� 569
13.6.3 Volatilization�� 569
13.6.4 Runoff and Leaching�� 570
13.7 Regulators�� 570
13.7.1 Effect of Electron Acceptors on Toxic Organic Degradation�� 570
13.7.2 Bacterial Groups�� 571
13.7.3 Effect of Soil Redox–pH Conditions on Degradation�� 572
13.7.4 Burial�� 577
13.8 Summary�� 577
Chapter 14 Soil and Floodwater Exchange Processes�� 579

14.2 Advective Flux�� 581
14.2.1 Advective Flux Processes�� 581
14.2.2 Measurement of Advective Flux�� 583
14.2.2.1 Seepage Meters�� 584
14.2.2.2 Piezometer�� 584
14.2.2.3 Salinity/Conductivity�� 584
14.2.2.4 Radium/Radon Isotopes�� 585
14.2.2.5 Dyes�� 585
14.3 Diffusive Flux�� 585
14.3.1 Diffusive Flux Processes�� 585
14.4 Bioturbation�� 591
14.4.1 Macrobenthos Communities�� 592
14.4.2 Benthic Invertebrates and Sediment–Water Interactions�� 593
14.5 Wind Mixing and Resuspension�� 594
14.6 Exchange of Dissolved Solutes Between Soil/Sediment
and the Water Column�� 595
14.6.1 Gradient-Based Measurements�� 596
14.6.2 Overlying Water Incubations�� 597
14.6.2.1 Benthic Chambers�� 597
14.6.2.2 Intact Core Flux�� 598
xiv Contents
14.7 Sediment Transport Processes��600

14.7.1 Sediment/Organic Matter Accretion in Wetlands�� 601
14.7.2 Measurement of Sedimentation or Accretion Rates�� 605
14.7.2.1 Filter Pad Traps��607
14.7.2.2 Artificial Marker Horizons��608
14.7.2.3 Sedimentation–Erosion Table��608
14.7.2.4 Beryllium-7 Dating�� 610
14.7.2.5 Lead-210 Dating�� 610
14.7.2.6 Cesium-137 Dating�� 613
14.7.2.7 Carbon-14 Dating�� 613
14.7.2.8 Application of Sediment Dating�� 614
14.8 Vegetative Flux/Detrital Export�� 614
14.9 Air–Water Exchange�� 616
14.10 Biogeochemical Regulation of Exchange Processes�� 617
14.11 Summary�� 619
Chapter 15 Coupled Biogeochemical Cycles: An Integrative Approach�� 621

15.2 Biotic Communities and Interactions�� 623
15.2.2 Periphyton�� 625
15.2.3 Vegetation�� 627
15.3 Coupled Biogeochemical Processes�� 628
15.3.1 Carbon�� 628
15.3.2 Nitrogen�� 631
15.3.3 Phosphorus�� 636
15.3.4 Sulfur�� 639
15.4 Ecological and Environmental Significance�� 642
15.4.1 Wetlands and Climate Change�� 642
15.4.2 Wetlands and Sea Level Rise��644
15.4.3 Wetlands and Water Quality�� 649
15.5 Summary�� 656
15.6 Future Directions and Perspectives�� 656
References�� 661
Index�� 704
Preface Preface Biogeochemistry
Wetland science has now emerged as a discipline where hydrologists, biogeochemists, pedologists,
ecologists, microbiologists, and scientists from various other disciplines are working individually
and together to improve our understanding of the functions and ecosystem services of wetlands.
The idea for this book came during the early 1980s when the first author began teaching a new
course entitled “Biogeochemistry of Wetlands” at the University of Florida. A similar course was
also taught at Louisiana State University during the early 1970s under the title of “Chemistry
of Flooded Soils,” and later in the 1980s the course title was changed to “Biogeochemistry of
Wetland Soils and Sediments.” Requests for a book were constant by the 1000+ students who took
this course before the first edition of the book was published in 2008. A similar interest to have
a wetland biogeochemistry reference book was also expressed by colleagues working with other
universities, governmental agencies, and other organizations.
The 2008 edition of the book primarily focused on the role of biogeochemistry as the key “oper-
ating system” that regulates the physical, chemical, and biological processes of elemental cycles
within a wetland, thereby affecting large-scale ecosystem services. We focused on “organic mat-
ter” as a hub of biogeochemistry and oxidation–reduction reactions as primary drivers of biogeo-
chemical processes. Wetlands are unique in that a range of soil–sediment conditions, from strongly
reducing (anaerobic) to oxidizing (aerobic), can be found at a range of spatial and temporal scales.
These environments include a wide range of systems such as tidal freshwater and salt marshes,
inland freshwater marshes and northern peat lands, swamp and bottomland forests, and ripar-
ian wetlands and estuaries. In certain topical areas, we relied heavily on the biogeochemistry of
aquatic systems because a limited amount of information was available on wetlands.
The first edition was written as both a reference and a text for a graduate-level course. Individuals
with an interest in environmental science, biology, chemistry, ecology, and environmental engi-
neering would also find this book useful. The book is highly cited by researchers from various
disciplines. The impact of soil redox processes on elemental cycling, biotransformation, and heavy
metal chemistry was emphasized. The book includes chapters dealing with terminology and elec-
trochemical properties describing basic biogeochemical processes that drive transformation pro-
cesses in wetlands. Detailed chapters included carbon, oxygen, nitrogen, phosphorus, sulfur, iron
and manganese, metals and metalloids, and toxic organics. In other chapters, we discussed plant
adaptation to wetland conditions, climate change, greenhouse gas emissions, and case studies
where extensive advancements have been made.
We wrote the second edition because the revisions to the first edition were long overdue. Our
revision was based on the feedback from colleagues and from the many University of Florida stu-
dents who used the first edition book in the “Biogeochemistry of Wetlands” course. As a result of
this feedback, many corrections and changes were made in the second edition. First and foremost,
we added a third author (Patrick Inglett) to the book to have an impact and thinking of the next
generation of wetland science experts. In the second edition, we added some new concepts and
helped streamline some of the topic areas for better application to emerging areas of research.
During revisions, we removed the last five chapters from the first edition and reincorporated some
of those key messages into other chapters. The second edition is also significantly improved with
the addition of many new references, special topic boxes to improve the teaching quality, and all
images are now in color.
The focus of the second edition was to provide an integrative approach to wetland biogeochemi-
cal cycles and the coupling and decoupling of macroelements (carbon, nitrogen, phosphorus, and
sulfur) as related to climate change, sea level rise, and water quality. We added a new synthesis
chapter that presents an integrative approach to coupled biogeochemical cycles and its environ-
mental and ecological significance. We sincerely hope the second edition provides state-of-the-art
xv
xvi Preface
scientific information involving wetland and aquatic systems biogeochemistry. As always, when
we wrote this book, we learned even more and made our best attempt to convey a contemporary
message on all aspects of wetland biogeochemistry. For those who read this book as a student, or as
an instructor, or others using it as reference material, we appreciate your comments and feedback
to help guide the next edition (planned in the next three to four years). We hope this book builds
on the first edition and continues to unite scientists of all disciplines in the study of wetlands and
serves as a foundation for building and inspiring the next generation of wetland professionals.
Acknowledgments Acknowledgments Biogeochemistry
We extend our sincere thanks to many of our colleagues who reviewed the content of the first edi-
tion, provided information to support the chapters, and helped to improve the quality of the book.
These include M. Clark, R. Corstanje, E. D’Angelo, W. F. DeBusk, M. Fisher, W. Hurt (deceased),
R. Gambrell, K. S. Inglett, J. Jawitz, T. Osborne, J. White, and many graduate students, postdoc-
toral fellows, and visiting scientists of the Wetland Biogeochemistry Laboratory at the University
of Florida. Many illustrations presented in this book were originally developed as part of a gradu-
ate course taught at the University of Florida, but the first two authors appreciate Patrick W. Inglett
for his efforts to create, update, and redraw the figures in both the first and second editions. All
three of us sincerely appreciate the contributions of those who came before us (on whose shoulders
we stand), as well as the creativity and hard work of the many students and post-docs we have
advised. Lastly, we gratefully acknowledge the support and love of our spouses (Sulochana Reddy,
Carole DeLaune [deceased], and Kanika Inglett) and families, without which this type of effort
simply would not be possible, or nearly as rewarding.
K. Ramesh Reddy
Institute of Food and Agricultural Sciences
School of Natural Resources and Environment &
Wetland Biogeochemistry Laboratory
Soil, Water and Ecosystem Sciences Department
Gainesville, Florida
Ronald D. DeLaune
Louisiana State University
Wetland Biogeochemistry Institute
Department of Oceanography and Coastal Sciences
School of the Coast and Environment
Baton Rouge, Louisiana
Patrick W. Inglett
Institute of Food and Agricultural Sciences
Wetland Biogeochemistry Laboratory
Soil, Water and Ecosystem Sciences Department
Gainesville, Florida
xvii
Authors Authors Biogeochemistry
K. Ramesh Reddy is a graduate research professor and director (2019 to present) of the School of
Natural Resources and Environment at the University of Florida (UF). Dr. Reddy served as chair
(2000–2018) of the UF Soil and Water Sciences Department. Dr. Reddy carried out research for
five decades on biogeochemical cycling of nutrients in natural and managed ecosystems as related
to water quality, carbon and nutrient sequestration, and greenhouse gas emission. Dr. Reddy taught
the “Biogeochemistry of Wetlands” course at UF for 40+ years. Dr. Reddy published 420+ refereed
journal articles and book chapters and edited five books. During his tenure at UF, Dr. Reddy served
as major advisor for 34 PhD and 20 MS students and served on 120+ graduate student committees.
Dr. Reddy has served on numerous advisory committees at state, national, and international levels.
Dr. Reddy’s select awards and honors include 1993: appointed as UF Graduate Research Profes-
sor (distinguished professorship); UF Doctoral Dissertation Advisory/Mentoring Award; Environ-
mental Quality Research Award, American Society of Agronomy; Soil Science Applied Research
Award, Soil Science Society of America; Fellow, American Association for the Advancement of
Science; Fellow–Soil Science Society of America; Fellow–American Society of Agronomy; 2012
Lifetime Achievement Award–INTECOL–Wetlands; 2016 National Wetlands Award-Research–
Environmental Law Institute, Washington DC; and 2016 Lifetime Achievement Award–Society of
Wetland Scientists. Dr. Reddy obtained a BS and MS from A.P. Agricultural University, India, and
a PhD (1976) from Louisiana State University.
Ronald D. DeLauneis an emeritus research professor in the Wetland Biogeochemistry Group,

Department of Oceanography and Coastal Sciences, School of the Coast and Environment,
Louisiana State University, Baton Rouge, Louisiana. His research interests include nutrient cycling
in agriculture and wetland ecosystems, role of wetlands in global biogeochemical cycles, wet-
land soil–plant interactions, coastal restoration and processes governing marsh stability, redox
chemistry of flooded soil and sediment, water quality of aquatic environments, greenhouse gas
emissions from wetlands, and the fate of organic pollutants and toxic heavy metals entering wet-
lands. Dr. DeLaune has served as associate editor or on the editorial board of numerous scientific
journals. He has authored more than 350 scientific papers in peer-reviewed journals. Dr. Delaune
is also a fellow in the Soil Science Society of America. Dr. DeLaune received his BS and MS from
Louisiana State University, and PhD from University of Wageningen, The Netherlands.
Patrick W. Inglettfirst became involved in wetlands when he worked in the Wetlands Protection
Section at the U.S. Environmental Protection Agency in Atlanta, Georgia. His interest was the sci-
ence behind the conservation and regulatory process for these unique ecosystems. He received his
BS in applied biology with a minor in earth and atmospheric sciences from the Georgia Institute
of Technology, and later, his MS and PhD from the University of Florida. Dr. Inglett is currently
a professor of biogeochemistry in the Soil and Water Sciences Department at the University of
Florida, where he also serves as the director of the Wetland Biogeochemistry Laboratory core
analytical and research facility. His research interests include nutrient cycling processes related to
ecosystem function and restoration, particularly aquatic systems. He specializes in nitrogen and
phosphorus limitation, enzymes, nitrogen fixation, temperature sensitivity of microbial processes,
greenhouse gas production, and the use of stable isotopes. For the last 15 years, Dr. Inglett has
taught both undergraduate and graduate courses in Environmental Biogeochemistry, Advanced
Biogeochemistry, and on occasion, the Wetlands Biogeochemistry course. He has advised more
than 50 graduate and undergraduate students, post-docs, and visiting scholars, and he is the author
of more than 60 journal articles and book chapters.
xix
1 Introduction
Wetlands are unique ecosystems located in areas with comparatively low elevation and a high
water table. Wetlands can include marshes, swamps, bogs, and similar areas that are poorly drained
and retain water during rainy periods. Globally, wetlands can be found in all climates, from tropi-
cal to tundra, with the exception of Antarctica. Approximately 6% of Earth’s land surface, which
equals approximately 1,200 million ha., is covered by wetlands (Ramsar Convention on Wetlands,
2018; Davidson and Finlayson, 2018). The United States alone accounts for approximately 10% of
the global wetlands, or 115 million ha. of wetlands, with Alaska representing approximately 62%
of the total wetland area. The conterminous United States accounts for approximately 44 mil-
lion ha., which includes 95% freshwater wetlands and 5% intertidal coastal wetlands (Dahl, 2011).
The Convention on Wetlands, signed in Ramsar, Iran, in 1971, is an intergovernmental treaty that
provides the framework for national action and international cooperation for the conservation and
wise use of wetlands and their resources. There are presently 158 contracting parties to the conven-
tion, with 1,723 wetland sites, totaling 160 million ha., designated for inclusion in the Ramsar List
of Wetlands of International Importance (http://www.ramsar.org/).
The value and function of wetlands are well recognized, as evidenced by national and
international policies to preserve wetland ecosystems. Why are wetlands so worthy of protec-
tion? Wetlands are complex ecosystems with functions driven by interactions of many physi-
cal, chemical, and biological processes. Wetlands are some of the most biologically productive
ecosystems on Earth; their productivity can exceed that of terrestrial and aquatic systems.
Wetlands not only serve to promote and sustain biota in many forms but also serve as living fil-
ters that process pollutants from terrestrial runoff and atmospheric deposition. Biodegradation
of organic compounds, elemental cycling, atmospheric exchange, hydrologic processing and
capacity, and plant response are controlled by the unique conditions found in the wetland envi-
ronment (Figure 1.1).
In the past, scientists often described wetland ecosystems on the basis of their disciplinary
bias, with an emphasis on one of the specialties such as hydrology, chemistry, wildlife, micro-
biology, or vegetation; however, no one discipline or specialization can describe these complex
processes. This was like the fabled group of blind men trying to describe an elephant by each
touching a different part, such as the side or the trunk or the tail. When the blind men compared
their notes, they were in complete disagreement in describing an elephant. The one who had
touched the elephant’s side claimed an elephant was like a wall; the man who had felt the tail
said an elephant was like a rope. It took the whole group of blind men to accurately describe the
elephant. Similarly, it has become very clear that no single discipline can adequately describe
a complex ecosystem such as wetlands. Instead, describing a wetland ecosystem requires an
interdisciplinary approach linking various specializations—biology, biogeochemistry, ecology,
environmental science, hydrology, and so on. Understanding must also draw on disciplines out-
side these fields. For example, much of the chemical and microbiological processes measured
in wetland soils is based on studies of saturated soils, which began around the turn of the 20th
century with research into nutrient behavior in paddy soils and processes measured in lake and
marine sediments.
In landscapes, wetlands typically occur between upland and aquatic ecosystems. Because
uplands are often the source of water to wetlands, components within runoff are also supplied
from uplands. In the absence of wetlands, contaminants added to or generated within upland areas
are directly transported into receiving aquatic ecosystems. Several physical, chemical, and bio-
logical processes functioning in the soil of uplands and wetlands are involved in regulating the
DOI: 10.1201/9780429155833-1 1
2 Biogeochemistry of Wetlands
Fertilizers, Animal Wastes

Biosolids, Wastewater
Uplands
Sink/source
Wetlands
Sink/source
Aquatic Systems
Sink/source
FIGURE 1.1 Linkages between uplands, wetlands, and aquatic systems.
fate (availability) of contaminants. For example, uplands and wetlands can serve as both “sinks,”
“sources,” and “transformers” for contaminants:
• Sink: Contaminants are transformed into biologically unavailable forms within the sys-
tem. For example, wetlands can convert nitrate to N2 gas through a biological reaction
called denitrification (this process is discussed in detail in Chapter 8).
• Source: Contaminants are transported from one ecosystem to another. For example,
uplands can serve as a “source” for suspended solids, nutrients, and other contaminants to
wetlands. Similarly, eutrophic wetlands can be a “source” of contaminants or nutrients to
adjacent aquatic systems such as streams, rivers, lakes, and estuaries.
• Transformer: Contaminants added to a wetland can also be transformed and released as
different or complexed compounds, or as new compounds to the aquatic ecosystem down-
stream. Because wetlands receive runoff from upland ecosystems, the changes in wetlands
can be used as an indicator of an upland ecosystem’s “health.”
Because wetland soils can serve as sinks, sources, and transformers of nutrients and other chemical
contaminants, they have a significant impact on water quality and ecosystem productivity.
The functions of wetlands in regulating nutrient and elemental storages and transformations are
captured in the study of biogeochemistry. Biogeochemistry is defined as the study of the exchange
or flux of materials between living and nonliving components of the biosphere [Box 1.1]. Like
the study of wetlands, biogeochemistry is also an interdisciplinary science, involving the interac-
tion of complex physical, chemical, and biological processes in various components of the ecosys-
tem, including the exchange of materials between biotic and abiotic. As defined, biogeochemistry
encompasses interactions from the smallest scale to the global scale encompassing the biosphere.
Wetland biogeochemistry, as the focus of this book, principally relates to small-scale exchanges
from the particle and microbial scale to the field scale. However, the cumulative impact of these
Introduction 3
small-scale processes on a landscape and on global reservoirs can be significant and will also be
addressed.
BOX 1.1 A BRIEF HISTORY OF BIOGEOCHEMISTRY

Contrary to most opinions, the idea of biogeochemistry or elemental cycling in earth systems
is not a very recent concept. You may be surprised to find that while the term biogeochemis-
try was first coined only in the early 20th century, the concept and foundation for this disci-
pline are found centuries earlier.
The roots of biogeochemistry began with the strong need for scientific achievement in
food production and agriculture. Many of the early scientists were also pioneers of science
itself, and nature, soils, and plants were often the subject matter for their basic physics and
chemical theories. As early as 1720, the derivation of materials for living biota were known to
come from gaseous and soil components, and the pioneer of chemistry, Lavoisier, correctly
found that according to conservation of mass, the gaseous material required for living plants
(carbon dioxide) was derived from the decomposition of plant biomass. In essence, this is the
first documented biogeochemical cycle. Later work by von Liebig and others properly related
the role of soil solution in supplying other water and particular or limiting nutrients to plant
growth. Liebig’s work was so important that his “law of the minimum” remains one of the
central concepts of nutrient limitation theory in biogeochemistry.
With the connection between plants and soils established, the concept of biogeochemis-
try gained more direction through the work of the founding fathers of soil science, includ-
ing Vasily Dokuchaev, who linked the composition and creation of soil materials with the
biota growing on them. Many of these scientists were interested in mineral composition and
diagenesis, so their studies centered on the relationship between biota living in ecosystems
with the mineral matter of the rocks and soils on which they were based. In 1875, Eduard
Suess coined the term “biosphere” and gave us the idea that soils and their associated biota
were really an intermediate in the flow of materials between the lithosphere, hydrosphere,
and atmosphere. These global cycling concepts were reinforced by advancements in geology,
including concepts of the rock cycle pioneered by James Hutton in the late 18th century, but
only gaining wide acceptance almost 100 years later. It was also at the end of the 19th century
that Walther Nernst established the thermodynamic principles of reduction and oxidation
chemistry leading to the creation of the Nernst equation.
Almost at the same time in the later part of the 19th century, important discoveries were
being made on a much, much smaller scale. These were the great advancements leading to
the birth of modern microbial ecology. After discoveries that microorganisms were involved
in the putrefaction of organic matter and fermentation, scientists like Winogradsky and
Beijerinck properly tied microbial identity and function to global elemental cycles with the
discovery of processes like nitrogen fixation, nitrification and denitrification, and sulfate
reduction. These process discoveries were pivotal in completing our understanding of the
cyclic nature of elements, and particularly that of nitrogen. Early work of Waksman, a bio-
chemist and microbiologist, expanded on the application of microbial processes with studies
on microorganisms and soil fertility, decomposition of plant and animal residues, formation
of humus, and the occurrence and role of bacteria in marine processes.
The discoveries of microbial ecology were earth shattering, leading scientists to begin
to see the cycles and spheres of the earth as having the ability to be sustainable or self-
regulating. In the early 1900s, a mineralogist and geochemist, Vladimir Vernadsky, saw
the importance of these types of processes as they related to the recycling of elements at the
(Continued)
(Continued)
Atmosphere
Biosphere
Hydrosphere Lithosphere
FIGURE 1.2 Conceptual diagram of the study of biogeochemistry as an interaction of the non-living
earth spheres (lithosphere, hydrosphere, and atmosphere) with life in the biosphere.
FIGURE 1.3 Vladimir Vernadsky, the father of modern biogeochemistry, used the early disciplines
of geochemistry and soil science to expand the concept of the biosphere and elemental cycling through
biota.
ecosystem and global scales. His theories expanded on the biosphere concept of Suess, where
he not only acknowledged that biota were “geologic force” but added that the biosphere was
in unity with the cycling of elements through the other geologic spheres (the lithosphere, the
atmosphere, and the hydrosphere). This idea is still at work today through concepts such as
Gaia, where the earth is viewed as a living organism.
Introduction 5
Because of his advancement of the biosphere concept and his original studies of element
budgets in ecosystems, Vernadsky is often credited as the father of modern biogeochemistry.
But it was not until a few more pieces of the puzzle were added that our modern view of bio-
geochemistry was truly realized. Following the acceptance of continental drift theory in the
1930s and 1940s, the final pieces of the modern biogeochemical puzzle were added with the
emergence of ecology in the 1950s and 1960s with scientists like Hutchinson and Redfield
and later Odum, who linked the populations of biota in ecosystems with their physical and
chemical environments.
It was also in that time that the study of oxidation-reduction coupled to biogeochemical
processes of paddy soils, lake and marine sediments, waterlogged, and poorly drained soils
began to be appreciated by early scientists in the 1930s to 1950s, including Sturgis, Pearsall,
Mortimer, Patrick, Ponnamperuma, and many others not included. The next generation of
scientists in the early 1970s (including the authors of this book) and their subsequent discov-
eries have expanded the discipline with major wetland biogeochemistry research programs
at several universities in the United States and in other countries. In the 1990s and through
today, we have seen the emergence of wetland and aquatic systems biogeochemistry as a
proper scientific discipline complete with dedicated journals, textbooks, and scientists will-
ing to label themselves as biogeochemists.
Many of the problems being addressed by biogeochemistry are large in nature (e.g., ocean
productivity, climate change), and some may hold an elitist view that biogeochemistry is only
defined at the global scale. However, as demonstrated in this brief history of the concept,
many scales are involved in the coupled cycling of elements, from microbes to continents.
All of these related disciplines and their discoveries are to be credited with developing this
notion to understand the interaction of life with the physical and chemical environments of
the Earth.
Biogeochemical cycles are influenced by various processes that result in exchange of materi-
als between two storage pools. The amount of a given constituent in these pools depends on its
residence time, which is simply the amount of that material in the reservoir divided by the rate
at which it is removed or added to the reservoir or the rate at which it is transformed. Living
components in wetland soils can act as exchange or cycling pools, with rapid turnover and flow
between the organisms in the pool and their immediate environment. The exchange or cycling
pool can encompass up to 20% of the total amount of a given compound of a system and turn over
rapidly, immobilizing and remobilizing compounds in a short time. Reservoir pools, which are
larger with slower turnover, provide long-term storage. The reservoir pool typically contains the
majority of a given compound in a system, is less reactive, and provides long-term storage. As an
example, when wetlands are used for wastewater treatment, designs that increase the percentage
of contaminants in the reservoir pool are more desirable because this provides long-term removal
of the contaminant.
Wetland biogeochemistry involves processes by which an element or a compound is trans-
formed within wetlands, including means by which various forms are interchanged between
the solid, liquid, and gaseous phases. Thus, the broader ecosystem biogeochemistry definition
is also applied to wetland biogeochemistry, which focuses on surface or near-surface processes
in wetlands that govern biogeochemical cycles, plant production, microbial transformations,
nutrient availability, pollutant removal, heavy metal chemistry, atmospheric exchange, and sedi-
ment transport. In this way, wetland biogeochemistry has its parentage in biology, soil science,
chemistry, and geology and now takes its place along with the bio-and geosciences in provid-
ing the knowledge needed to develop solutions to the challenges faced by wetland and global
ecosystems.
Biogeochemical processes in wetlands involve a host of complex microbial communities,

periphyton, and vegetation, and interaction and mutual dependency among these communities.
Biogeochemical processes also quantify the exchange and transport of elements or compounds
within wetlands, including those to other systems (e.g., the atmosphere). Wetland biogeochemical
processes provide overviews of the environmental factors and alterations in wetlands that control
or affect functioning at the local, regional, and global level and can have both positive and negative
feedbacks. Nitrogen and carbon cycles are good examples of biogeochemical processes that affect
processes at the local level (e.g., plant growth and soil accretion, regional level (water quality), and
the global level (greenhouse gas emissions and carbon storage). Wetland biogeochemical cycles
can have global significance as follows (Figure 1.4):
• Eutrophication of oligotrophic wetlands resulting from increased nutrient loads can

enhance primary productivity.
• High nutrient loads can affect species composition and food webs, leading to a dominance
of fast-growing, high-reproduction organisms.
• High primary productivity can result in increased rates of organic matter accumulation,
providing a sink for carbon (increased carbon sequestration).
• High rates of carbon accumulation can enhance microbial activities in soil and the water
column, leading to increased nutrient regeneration and biotic growth.
• Increased rates of microbial activities can increase the production of greenhouse gases,
and increased levels of greenhouse gases can have a negative impact on climate.
As we walk through a wetland, we all admire beautiful plants, flowers, birds and other wildlife,
and flowing water, but we rarely think about the “living soil” under our feet. The biogeochemical
Physical
Processes
Chemical Biological
Organic Matter Processes Processes
N C P
Climate
S Change
Carbon Eutrophication
Sequestration
FIGURE 1.4 Linkages between physical, chemical, and biological processes and global-scale processes in
the biosphere.
Introduction 7
processes in the soil control many functions and ecosystem services provided by wetlands. This
is similar to the “brain” orchestrating the many functions of the human body. The type of biogeo-
chemical transformation occurring in wetlands, in contrast to upland systems, is strongly gov-
erned by hydrology. Transformations that occur in wetlands involve both anaerobic and aerobic
processes. The chapters presented in this book focus on the role excess water and soil oxidation–
reduction processes play in elemental cycling, heavy metal transformation, wetland plant response,
and toxic organic transformation. The “hub” for biogeochemistry is organic matter, which contains
macroelements including carbon, nitrogen, phosphorus, potassium, and sulfur (Figure 1.5). The
cycling of these macroelements in the aerobic and anaerobic zones of the soil and water column is
a primary driver supporting ecosystem processes and is far more interesting and important than
any other constituent of wetlands.
Nutrients such as nitrogen, phosphorus, and sulfur are primary components of soil organic mat-
ter, and the cycling of these nutrients is always coupled with carbon cycling. The rate and extent
of many of these reactions in the soil and the water column, involving carbon, nitrogen, phospho-
rus, and sulfur, are mediated by microbial communities and associated physicochemical reactions.
These cycles are coupled, and there is considerable mutual dependency of one cycle on another
(feedbacks and controls) or one organism on another (microbes, algae, and vegetation). There is a
strong linkage between biogeochemical processes and biotic communities (vegetation, algae, and
microbes). These coupled cycles operate at different spatial and temporal scales from the molecular
to landscape level.
As you can see, wetlands and their biogeochemical processes are key drivers of several ecosys-
tem functions associated with wetland values (e.g., water quality improvement through denitrifi-
cation and long-term nutrient or carbon storage in the organic matter; Figure 1.5). Wetlands are
Plant Community Loading Hydroperiod
Carbon (productivity)
S P
Metals
Stable Organic Matter

(P Accretion/Stability)
(Accretion/Stability)
FIGURE 1.5 Relationship between coupled biogeochemical cycles and organic matter accretion in wetlands.
atmospheric sources of carbon dioxide, methane, and nitrous oxide, and due to flooded or reducing
conditions, wetlands also limit organic matter turnover to serve as important global carbon sinks.
Thus, biogeochemical processes in wetlands are also important in global issues, such as global
warming, carbon sequestration, and eutrophication and water quality. For these reasons, knowl-
edge of wetland biogeochemical processes is useful both for understanding the ecology of these
systems leading to their preservation and to help us manage and predict the environmental fate and
transport of elements and compounds from natural or anthropogenic sources. In both ways, the
study of wetland biogeochemistry from the micro-to the macro-scale is of critical importance for
the challenging issues we face today.
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Biogeochemistry of Wetlands: Science and Applications

Book · October 2022

Konda Ramesh Reddy Ronald D Delaune

SEE PROFILE SEE PROFILE

The user has requested enhancement of the downloaded file.

• Flooded soil and sediment characteristics

K. Ramesh Reddy, Ronald D. DeLaune and

Chapter 2 Basic Concepts and Terminology��������������������������������������������������������������������������������9

Chapter 3 Biogeochemical Characteristics��������������������������������������������������������������������������������� 33

3.3 Wetland Hydrology����������������������������������������������������������������������������������������� 41

Chapter 7 Adaptation of Plants to Soil Anaerobiosis��������������������������������������������������������������� 239

8.3.5 Inorganic Forms of Nitrogen������������������������������������������������������������ 285

10.4 Reduction of Iron and Manganese���������������������������������������������������������������� 435

11.10 Biogenic Emission of Reduced Sulfur Gases����������������������������������������������� 496

Chapter 13 Toxic Organic Compounds�������������������������������������������������������������������������������������� 543

Chapter 14 Soil and Floodwater Exchange Processes���������������������������������������������������������������� 579

14.7 Sediment Transport Processes����������������������������������������������������������������������600

Chapter 15 Coupled Biogeochemical Cycles: An Integrative Approach����������������������������������� 621

Ronald D. DeLauneis an emeritus research professor in the Wetland Biogeochemistry Group,

Fertilizers, Animal Wastes

FIGURE 1.1 Linkages between uplands, wetlands, and aquatic systems.

BOX 1.1 A BRIEF HISTORY OF BIOGEOCHEMISTRY

Biogeochemical processes in wetlands involve a host of complex microbial communities,

• Eutrophication of oligotrophic wetlands resulting from increased nutrient loads can

Plant Community Loading Hydroperiod

Stable Organic Matter

Adaptation of Plants to Soil Anaerobiosis

Toxic Organic Compounds

Soil and Floodwater Exchange Processes

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Chapter 2 Basic Concepts and Terminology��9

Chapter 3 Biogeochemical Characteristics�� 33

3.3 Wetland Hydrology�� 41

Chapter 7 Adaptation of Plants to Soil Anaerobiosis�� 239

8.3.5 Inorganic Forms of Nitrogen�� 285

10.4 Reduction of Iron and Manganese�� 435

11.10 Biogenic Emission of Reduced Sulfur Gases�� 496

Chapter 13 Toxic Organic Compounds�� 543

Chapter 14 Soil and Floodwater Exchange Processes�� 579

14.7 Sediment Transport Processes��600

Chapter 15 Coupled Biogeochemical Cycles: An Integrative Approach�� 621