Nitrogen in the Marine Environment

Dedication Page......Page 2
A Timely Book for Interesting Times......Page 3
Preface to Second Edition......Page 4
Acknowledgements......Page 6
Contributors......Page 7
Copyright Page......Page 13
Introduction......Page 14
The electronic diversity of nitrogen......Page 17
The major reactions......Page 19
Inventories and residence times......Page 24
Global mean profiles......Page 25
Surface distribution......Page 27
Interior ocean distribution......Page 29
Distribution......Page 33
New versus regenerated production......Page 35
Nitrite......Page 37
Nitrogen*: The imprint of N2-fixation and denitrification......Page 39
Oceanic nitrogen budget......Page 45
Global nitrogen budget......Page 48
Controls on the marine fixed nitrogen content......Page 50
The marine nitrogen cycle, carbon cycle, and climate......Page 53
The anthropogenic perturbation......Page 54
References......Page 56
Gaseous Nitrogen Compounds (NO, N2O, N2, NH3) in the Ocean......Page 64
Climatic and biogeochemical relevance......Page 65
Unicellular cyanobacterial N2 fixers......Page 66
The role of the ocean for the global budget of atmospheric NOx......Page 67
Climatic and biogeochemical relevance......Page 68
Particle export and sub-euphotic zone remineralization of nitrate......Page 734
Water-column denitrification......Page 284
N2O in coastal areas......Page 75
Sponge-microbe specificity and phylogenetic diversity......Page 78
Isotope studies......Page 79
Distribution of nitrification in water and sediments......Page 235
Climatic and biogeochemical relevance......Page 84
N2 in the ocean......Page 85
Climatic and biogeochemical relevance......Page 88
The NH3/NH4+ equilibrium in water......Page 89
NH3 in the ocean......Page 92
The ocean as a source or sink of atmospheric NH3......Page 93
The role of oceanic emissions in the global budget of atmospheric NH3......Page 94
Outlook......Page 95
Nitrogen fixation......Page 902
Note added to proof......Page 96
References......Page 97
Chemical Composition of Marine Dissolved Organic Nitrogen......Page 108
Introduction......Page 521
Definitions of Dissolved Organic Nitrogen......Page 109
Solid phase extraction......Page 110
Coupling of nitrification, denitrification, dissimilatory nitrate reduction to ammonium, and anammox......Page 111
Bulk Chemical Composition of High Molecular Weight Dissolved Organic Nitrogen......Page 112
Adaptive strategies. Oxygen consumption-The Mehler reaction......Page 1103
Nitrogen requirements: Effects of food quantity and quality......Page 115
A nitrogen budget for the baltic sea......Page 117
Gaseous extraction......Page 1354
Proteins......Page 120
Amino acid composition......Page 121
The enantiomeric signature of amino acids......Page 125
Macroalgae as Indicators of Sources and Magnitude of Nitrogen Supply......Page 128
Isolation of proteins from HMWDOM......Page 131
Monosaccharide composition......Page 133
Molecular reconstruction of nitrogen in HMWDOM......Page 134
Sources......Page 138
Corals......Page 419
Residence time of DOM components......Page 145
References......Page 146
Introduction......Page 154
Benthic Nitrogen Fixation......Page 155
Bioavailability of Nitrogen Compounds......Page 156
Kinetic fractionation......Page 1278
Silica to nitrogen ratios energetics......Page 161
Microbial mats......Page 162
Rocky intertidal......Page 928
Pelagic Nitrogen Fixation......Page 165
Aeolian iron and stratification of the surface mixed layer in the future......Page 1548
What happens to all that fixed nitrogen?......Page 168
Geochemical constraints to estimate nitrification rates......Page 169
Recycling......Page 170
Current Challenges and Opportunities......Page 1292
Nitrogen fixation......Page 173
Amino acids......Page 1360
Trace metals......Page 174
Phosphorus limitation......Page 177
Detritus and marine snow......Page 420
Biogeochemical Significance of Marine Nitrogen Fixation......Page 181
Directly assessing nitrogen fixation......Page 185
Diazotroph densities......Page 189
Comparison of geochemical vs. direct estimates......Page 192
Summary and Future Directions......Page 194
References......Page 197
Nitrification in Marine Systems......Page 212
Introduction......Page 1074
Micro-nutrient iron......Page 214
Seasonal variations of surface nitrate......Page 220
Anaerobic nitrification......Page 222
Assessment of in situ Nitrogen-Status......Page 224
Methods for measuring nitrification rates in water and sediments......Page 229
N2-fixation......Page 1468
Isotopic incubation methods......Page 231
Variability in biological and dissolved nitrogen:phosphorus stoichiometry......Page 795
Particulate and sedimentary organic nitrogen......Page 244
Environmental Variables Affecting Nitrification Rates and Distributions......Page 247
Effects of shifting autotrophic dominance on nitrogen cycle processes......Page 896
Salinity......Page 248
Case Studies of Specific Enzymes......Page 249
Nitrogen uptake kinetics......Page 250
Substrate concentration......Page 253
Oxygen concentration......Page 254
Nitrification and Methane Oxidation......Page 256
Future Directions......Page 257
References......Page 261
Principles and Practice......Page 275
Canonical or respiratory denitrification......Page 277
Anammox......Page 278
Dissimilatory nitrate reduction......Page 281
Environmental controls on denitrification......Page 282
Rates of water-column denitrification......Page 286
Sedimentary denitrification......Page 290
Conversion to gaseous forms......Page 1356
Rates of sedimentary denitrification......Page 293
Sites and rates of anammox......Page 296
Isotopic Consequences of Denitrification......Page 299
Denitrification and the Marine Combined Nitrogen Budget......Page 301
Acknowledgements......Page 304
Introduction......Page 314
Re-Evaluation of Nitrogen Limitation and New Production in the Sea......Page 333
NH4+......Page 338
Nitrate......Page 1282
Patterns......Page 490
N2......Page 344
Bulk organic nitrogen dissolved......Page 346
Urea......Page 347
Comparisons with Other Enclosed Seas......Page 703
Recorders of the Past Nitrogen Cycle......Page 1355
Other organic compounds......Page 355
Nutrient/nutrient interactions......Page 356
Environmental interactions......Page 358
Ocean delta15Nnitrate through time......Page 1212
Pathways of Nitrogen Uptake and Assimilation......Page 361
Conclusions......Page 1295
The role of benthic suspension feeders in reducing eutrophication potential......Page 900
Regulation of Nitrogen Uptake and Assimilation......Page 367
Whole genome analyses......Page 369
What Does the Future Hold?......Page 370
References......Page 372
Nitrogen Regeneration......Page 396
Inorganic......Page 397
Sources of iron......Page 1498
Nitrate (NO3-) and nitrite (NO2-) analysis......Page 398
Animals and food webs......Page 399
Programmed cell death......Page 402
Mediated release......Page 403
Release by different phytoplankton groups......Page 404
N2 fixers......Page 405
Bacteria......Page 407
Viruses and other parasites......Page 409
Zooplankton......Page 411
Release by different zooplankton groups......Page 412
Grazing, bacterivory, and viral lysis in marine sediments......Page 1116
Nekton......Page 417
Sediments......Page 421
Variables that affect sediment nitrogen regeneration......Page 430
Sources of regenerated nitrogen associated with sediments......Page 431
Methods to Measure Nitrogen Regeneration......Page 434
NH4+ regeneration versus DON release......Page 436
Variables that affect NH4+ regeneration rates......Page 439
DON release versus nitrogen uptake......Page 448
Variables that affect DON release rates......Page 457
Individual organic compounds......Page 458
Linking DON release with DOC flux......Page 459
Recommendations for Future Research......Page 460
References......Page 461
Land-Based Nitrogen Sources and Their Delivery to Coastal Systems......Page 479
Spatial Patterns in Amount and Form of River Nitrogen Export......Page 481
Nitrate and oxygen......Page 641
Patterns......Page 483
Dissolved Inorganic Nitrogen Uptake in Upwelling Areas......Page 489
Long-term trends......Page 494
Interannual variability......Page 496
Seasonal variability......Page 497
High intensity, low frequency events......Page 501
Effects on Nitrogen Export of Long-Term Human Modification of Discharge......Page 502
Groundwater......Page 503
Atmospheric Deposition Directly to Coastal Waters......Page 505
Summary and Future Directions......Page 507
References......Page 513
Photochemical Production of Inorganic Nitrogen......Page 522
Photochemical Transformations of Organic Compounds and Effects on Bioavailability......Page 530
References......Page 535
Nitrogen and Marine Eutrophication......Page 538
The Evidence for the Role of Nitrogen in Marine Eutrophication......Page 543
Nutrient, Physical and Climatic Controls of Marine Eutrophication......Page 549
Is Nitrogen Nitrogen? Roles of Different Nitrogen Sources in Marine Eutrophication......Page 554
The Role of Nitrogen in Relation to other Nutrients......Page 557
Human development of the coastal landscape......Page 561
Agricultural activities......Page 563
The Future and Nitrogen Management......Page 566
References......Page 679
References......Page 568
Introduction......Page 577
Environmental Factors Regulating Nitrogen Uptake......Page 580
Nitrogen concentration effects......Page 585
Nitrogen substrate interactions......Page 587
Internal stabilizing feedbacks on the marine nitrogen inventory......Page 1499
Irradiance effects......Page 592
Organic complexation and redox chemistry......Page 1598
Summary......Page 595
Acknowledgements......Page 1486
References......Page 596
Introduction......Page 605
Experimental Design......Page 1198
Previous examinations of nitrogen distributions......Page 817
Dissolved organic nitrogen......Page 610
Physical supply of nitrate......Page 617
Geochemical investigations of nitrogen fixation......Page 620
Atmospheric deposition and possible contribution to excess nitrate......Page 625
Denitrification......Page 627
Effects on body nitrogen content......Page 1157
Is the Atlantic a Source of Nitrogen to the Global Ocean?......Page 628
Phosphorus versus nitrogen as proximate and ultimate limiting nutrients......Page 1639
References......Page 631
Introduction and Background......Page 639
General principles of nutrient analysis......Page 1222
Bacteria......Page 649
Denitrification......Page 653
Intermediate nepheloid layer and quantitative significance of denitrification in remineralization......Page 659
Nitrous oxide......Page 661
Nitrogen fixation......Page 665
Implications for the global carbon cycle......Page 666
Derivatization and chromatography......Page 672
Nitrogen Budgets......Page 676
Concluding Remarks......Page 677
Nitrogen in Inland Seas......Page 690
Introduction......Page 1135
The Model Inland Sea-The Baltic Sea......Page 691
Nitrogen in surface and deep water-Concentrations and pools......Page 693
Oxygenation of the early atmosphere: The influence of shelf area and nitrogen fixation rates......Page 694
Molecular Characterization of Nitrogen Assimilatory Pathways......Page 1316
Invertebrate/symbioses......Page 698
Ammonification and regeneration......Page 966
Conclusions and Future Research Prospects and Needs......Page 706
Acknowledgements......Page 707
References......Page 1373
The Nitrogen Cycle in the North Pacific Trades Biome: An Evolving Paradigm......Page 712
The North Pacific trades biome......Page 716
Evolutionary constraints for nitrogen fixation and adaptive strategies......Page 721
Modeling new production, NO3, and NH4......Page 1350
Nitrogen assimilation and particulate nitrogen production......Page 730
Selected Trades Biome Ecosystem Processes......Page 738
Plant and sediment stocks......Page 1000
Structure and dynamics of the primary and secondary nitrite maximum layers......Page 741
New production by nitrogen fixation......Page 746
Aperiodic delivery of nutrients and consequences for ecosystem metabolism......Page 761
Epilogue......Page 766
References......Page 767
Introduction......Page 777
Inputs and Concentrations of Dissolved Nitrogen......Page 779
Historical context for the microbial loop......Page 1100
Characteristic NO3 concentrations in different upwelling regions......Page 780
NO3 uptake and new production......Page 783
NH4 uptake and regenerated production......Page 786
Phytoplankton Functional Groups and Nitrogen Assimilation......Page 789
Physiological Adaptation of Nitrogen Assimilation......Page 790
Kinetics of NO3 and NH4 uptake......Page 792
Outputs......Page 794
Ecosystem models of nitrogen flux......Page 796
Local variations......Page 798
Internal pools......Page 1362
Molecular Approach to Studying Nitrogen Assimilation......Page 800
Conclusions and Directions......Page 802
Estuaries......Page 814
Recent Advances in Molecular Methods for Studying Uncultivated Marine Microorganisms Active in Nitrogen Cycling......Page 1303
Comparative examination of nitrogen distributions......Page 818
Nitrogen-Fixation, Nitrogenase, and Oxygen. Evolutionary Constraints, Adaptations of Cellular Pathways and Organisms, Feedbacks to Global Nitrogen, Carbon, Oxygen Cycles......Page 823
Nitrogen effects on seagrass community structure and distribution......Page 1043
Inputs of nitrogen to estuaries......Page 826
Approaches to enzyme activity measurements......Page 1393
Export of nitrogen to downstream systems......Page 837
Ecosystem metabolism: Interactions with nitrogen cycling......Page 839
Sediment-water Solute Fluxes......Page 842
Overview of flux magnitude......Page 843
Spatial patterns of flux......Page 845
Regulation of sediment-water fluxes......Page 846
Modern nitrogen-cycle model structures......Page 1480
Deep nitrate......Page 1507
Statistical models of nitrogen and production......Page 850
Nutrient limitation......Page 853
Benthic autotrophs and nitrogen......Page 854
Summary and Future Directions......Page 1062
Effects of nitrogen enrichment on secondary production......Page 856
Effects of estuarine animals on nitrogen-budgets and cycles......Page 858
Summary and Future Directions......Page 859
References......Page 861
Nitrogen Cycling in Coastal Sediments......Page 872
The Sediment Nitrogen Cycle: Overview......Page 873
Iron limited regimes......Page 1569
Ammonium regeneration by mineralization......Page 877
Interactions between the nitrogen and sulfur cycles......Page 884
Regulation of nitrogen cycle processes by oxygen......Page 885
Advective and diffusive transport through sediments......Page 887
Assays......Page 888
The influence of salinity on nitrogen cycle processes......Page 890
Biotic Factors Influencing Sediment Nitrogen Transformations......Page 891
Scaling activity......Page 1398
The influence of benthic macrophytes on nitrogen cycle processes......Page 892
Steady state mean......Page 897
Emission spectrometers......Page 1363
Transfer of benthic microalgal-derived carbon and nitrogen to higher trophic levels......Page 901
Glacial-interglacial inventory changes......Page 903
Growth and reproduction......Page 1177
Anammox......Page 905
Denitrification......Page 1370
References......Page 978
Macroalgal-Dominated Ecosystems......Page 921
Introduction......Page 1302
Basic definitions......Page 922
Coral reefs......Page 924
Rock reefs......Page 926
Nitrogen Limitation of Marine Macroalgae: Evidence and Approaches......Page 930
Sources of nitrogen to macroalgae......Page 932
Macroalgae as a sink of nutrients: nitrogen uptake......Page 934
Macroalgae as a sink of nutrients: nitrogen storage and retention......Page 936
Macroalgae as a source of nitrogen: turnover rates......Page 937
Knowledge Gaps and Future Directions......Page 941
Introduction......Page 952
New developments......Page 954
Nitrogen Cycle Processes......Page 955
Nitrification......Page 960
Nitrogen acquisition and uptake......Page 961
Microbial populations......Page 965
Nitrogen Perturbations to Reefs......Page 969
Surface-bound phosphorus......Page 970
Algal increases and grazer decreases......Page 971
Effects of runoff and sedimentation......Page 972
Analytical advances......Page 975
Introduction......Page 993
Background......Page 994
Plant production and biomass......Page 995
Marsh plant community structure......Page 997
Ammonium......Page 998
Nitrogen budgets for salt marshes......Page 999
Speciation of dissolved phosphorus......Page 1637
Denitrification......Page 1006
Ammonification......Page 1019
Ammonium Uptake......Page 1020
Marsh macrophyte nitrogen dynamics......Page 1021
Summary of salt marsh nitrogen fluxes......Page 1022
Eutrophication and Management......Page 1025
Role of Salt Marshes in Global Nitrogen Balance......Page 1026
Summary and Future Research Directions......Page 1027
References......Page 1261
Seagrass Habitats......Page 1039
Nitrogen Inputs to Seagrass Ecosystems......Page 1041
Viruses as a member of the microbial loop......Page 1042
Total dissolved nitrogen/dissolved organic nitrogen analysis......Page 1044
Uptake......Page 1045
Turnover in plant biomass......Page 1048
Fate of Assimilated Nitrogen......Page 1049
Mineralization and burial......Page 1050
Assay conditions......Page 1397
Litter export......Page 1053
Dissolved organic nitrogen......Page 1054
Denitrification......Page 1058
Ascidian-prochloron symbioses......Page 1060
Are Seagrass Meadows Sources or Sinks of Nitrogen?......Page 1061
General Aspects of Cyanobacterial Nitrogen Metabolism......Page 1075
Inventory changes......Page 1077
Unicellular non-N2-fixers......Page 1081
Regulation of Nitrogen Metabolism......Page 1084
Genomics......Page 1087
Cyanobacteria and the Nitrogen-Cycle......Page 1089
Future Perspectives......Page 1091
References......Page 1092
Importance of enzyme-mediated reactions......Page 1097
Components of the microbial loop......Page 1101
Carbon:nitrogen ratios in bacteria......Page 1106
Distribution of Bacterial Secondary Production in the Marine Environment......Page 1107
Causes and consequences of silica to nitrogen ratios changes......Page 1603
Harmful algal blooms......Page 1605
Virus abundance in marine waters and sediments......Page 1110
Rates of viral mortality and nitrogen release......Page 1112
Lysogeny and horizontal gene transfer......Page 1114
Marine Sediments Compared to the Water Column......Page 1115
Future Directions of Microbial Loop Research......Page 1117
Temperature......Page 1118
Viral and autocatalysed mortality of key nitrogen-processing organisms......Page 1119
Conclusion: A New Microbial View-the Gene Loop......Page 1120
References......Page 1121
Phytoplankton......Page 1137
Zooplankton......Page 1139
Diatoms versus coccolithophores......Page 1596
Effects on feeding and growth rates......Page 1155
Effects on egg production and hatching success......Page 1156
Response to variable food quality......Page 1158
Egestion......Page 1159
Sloppy feeding......Page 1163
Assimilation......Page 1164
Inorganic......Page 1165
Organic......Page 1171
Factors affecting excretion rates......Page 1175
The delta15N of sedimentary nitrogen......Page 1510
Other miscellaneous environmental factors......Page 1176
Conclusions and Future Directions......Page 1180
References......Page 1182
Introduction......Page 1197
Cultivation, transmission, and cell divisions......Page 1200
Specificity and symbiont phylogenetic diversity......Page 1201
Host-symbiont interactions......Page 1202
Geographical distribution and cell abundances......Page 1204
Nitrogen fixation......Page 1205
Organic extractions......Page 1208
Nitrification in sponges......Page 1209
Implications......Page 1211
The sponge-phototroph nitrogen trap......Page 1213
References......Page 1214
Analytical Methods for the Study of Nitrogen......Page 1219
Introduction......Page 1220
Collection and Storage of Samples......Page 1221
Ammonium (NH4+) analysis......Page 1223
Urea......Page 1228
Nitrate distribution and the nutrient-rich surface regions......Page 1501
Humic substances......Page 1229
Amino acids......Page 1230
Hydrolysis methods......Page 1231
Chiral (d/l) amino acids......Page 1233
Colorimetric methods for bulk protein......Page 1234
Specific protein analysis......Page 1235
Amino sugars......Page 1236
Nucleic acids......Page 1237
Bulk Organic Nitrogen Characterization......Page 1238
Ultrafiltration......Page 1239
Hydrophobic resin isolations......Page 1241
Dialysis......Page 1242
Looking backward......Page 1522
Basic principles......Page 1244
NH4+......Page 1245
DON......Page 1246
Particulate nitrogen......Page 1247
Substrate additions......Page 1248
Uptake rate calculations......Page 1249
DON release......Page 1251
Nitrification......Page 1253
Denitrification......Page 1254
N2 fixation......Page 1255
Reference materials......Page 1256
Hobbled by hydrolysis? Chipping away at the black box......Page 1257
DON Geomolecules? Exploring the nature of "humic" DON......Page 1258
Higher order of information: Proteomics in DON......Page 1259
Rates-Who is doing what?......Page 1260
Introduction......Page 1276
Isotopic fractionation and oceanic nitrogen......Page 1280
Particulate organic matter......Page 1283
Analytic Considerations......Page 1285
Blank corrections: Theory......Page 1286
Measurement of sample and blank size......Page 1287
Ecosystem Level Patterns in delta15N......Page 1288
Terrigenous inputs......Page 1289
Nitrogen sources for pelagic primary production......Page 1290
References......Page 1296
The Power of Genomics to Reveal Metabolic Potential......Page 1309
Nitrogen as a Nutrient......Page 1312
Identifying Microorganisms Involved in Nitrogen Uptake......Page 1313
Particles......Page 1351
Assimilation of DON......Page 1321
Identifying nitrogen limitation......Page 1323
Molecular Biological Tools for Probing the Use of Nitrogenous Compounds for Cellular Energy......Page 1324
Nitrogen fluxes and ocean circulation......Page 1476
Ammonia oxidation and nitrification......Page 1326
Summary and Perspective......Page 1328
References......Page 1329
Introduction......Page 1344
Initial planning......Page 1348
Incubation protocols......Page 1349
Gaseous nitrogen-N2O and N2......Page 1357
Urea......Page 1359
Interpreting enzyme measurements......Page 1400
Mass spectrometers......Page 1364
Basic equations......Page 1365
Isotope dilution......Page 1367
Comparisons between 15N rates and other methods......Page 1369
Future Directions......Page 1371
Enzymes and Nitrogen Cycling......Page 1384
Nitrogen acquisition......Page 1389
Incorporation of nitrogen into macromolecules and secondary compounds......Page 1390
Turnover and maintenance of internal nitrogen......Page 1391
Enzyme kinetic measurements......Page 1399
Overview......Page 1402
Assay techniques......Page 1403
Observations and Interpretations......Page 1512
Overview......Page 1405
Enzyme regulation and interpretation of activity assays......Page 1406
Looking forward......Page 1523
Information gained from assays and future potential......Page 1408
Ammonium assimilation pathways......Page 1409
Assay techniques......Page 1410
Enzyme regulation and interpretation of activity assays......Page 1411
Assay techniques......Page 1413
Overview......Page 1414
Overview......Page 1415
Enzyme regulation and interpretation of activity assays......Page 1416
Overview......Page 1417
Assay techniques......Page 1418
Enzyme regulation and interpretation of activity assays......Page 1420
Overview......Page 1421
Information gained from assays and future potential......Page 1422
Assay techniques......Page 1423
Assay techniques......Page 1424
Assay techniques......Page 1425
Summary......Page 1426
References......Page 1427
Ocean Nitrogen Cycle Modeling......Page 1444
Evolution of Nitrogen-cycle Modeling......Page 1446
Modeling phytoplankton growth and cell quota......Page 1455
Loss terms, remineralization and the microbial loop......Page 1461
Modeling export......Page 1467
Denitrification......Page 1472
Mesoscale processes......Page 1479
Future Challenges and Conclusions......Page 1482
References......Page 1487
Introduction......Page 1495
Potential Changes in the Marine Nitrogen Cycle During the Past 2 Million Years......Page 1497
N2 fixation self-limiting feedback......Page 1500
Inputs......Page 1503
Outputs......Page 1505
Internal cycling......Page 1506
Application to the past......Page 1508
Sedimentary nitrogen isotope records......Page 1509
Glacial-interglacial changes in nutrient-rich regions......Page 1519
Connections......Page 1521
References......Page 1524
Nitrogen in the preoxygenated world/oceans......Page 1534
Biological feedbacks to the global nitrogen, carbon, and oxygen cycles......Page 1546
Sensitivity of the Nitrogen, Carbon, and Oxygen Cycles to Climate Change......Page 1547
Ocean circulation and the global ocean cycles of nitrogen, oxygen and carbon......Page 1550
Summary and Conclusions......Page 1553
Acknowledgements......Page 1554
Introduction......Page 1561
Nitrogen Budget Method......Page 1568
Nutrient transport kinetics......Page 1571
Distribution of Nitrogen Fluxes in Rivers by Continent and Ocean Basin......Page 1573
Development of Policy and Management Strategies for Reducing Coastal Nitrogen Pollution......Page 1576
References......Page 1580
Silicon:Nitrogen Interactions in the Marine Environment......Page 1584
Molecular ecology of diatoms......Page 1586
The southern ocean......Page 1601
Pseudo-nitzschia: A case study......Page 1606
Impacts on higher trophic levels......Page 1607
Conclusions......Page 1608
References......Page 1609
Linking the Oceanic Biogeochemistry of Iron and Phosphorus with the Marine Nitrogen Cycle......Page 1622
The Marine Iron Cycle......Page 1623
Iron limitation and marine food webs......Page 1625
Iron export and residence time......Page 1628
Dissolved and particulate iron pools......Page 1630
Iron and nitrogen acquisition......Page 1633
The Ocean Phosphorus Cycle......Page 1635
Phosphosrus and Iron as Alternative or Co-limiting Controls on N2 Fixation......Page 1642
Conclusions and Future Directions......Page 1648
References......Page 1649
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