Nitrogen cycle of the open ocean: from genes to ecosystems.

The marine nitrogen (N) cycle controls the productivity of the oceans. This cycle is driven by complex biogeochemical transformations, including nitrogen fixation, denitrification, and assimilation and anaerobic ammonia oxidation, mediated by microorganisms. New processes and organisms continue to be discovered, complicating the already complex picture of oceanic N cycling. Genomics research has uncovered the diversity of nitrogen metabolism strategies in phytoplankton and bacterioplankton. The elemental ratios of nutrients in biological material are more flexible than previously believed, with implications for vertical export of carbon and associated nutrients to the deep ocean. Estimates of nitrogen fixation and denitrification continue to be modified, and anaerobic ammonia oxidation has been identified as a new process involved in denitrification in oxygen minimum zones. The nitrogen cycle in the oceans is an integral feature of the function of ocean ecosystems and will be a central player in how oceans respond during global environmental change.

[1]  E. Carpenter,et al.  Marine Cyanobacterial Symbioses , 2022, Biology and Environment: Proceedings of the Royal Irish Academy.

[2]  Nicole L. Goebel,et al.  Abundance and distribution of major groups of diazotrophic cyanobacteria and their potential contribution to N₂ fixation in the tropical Atlantic Ocean. , 2010, Environmental microbiology.

[3]  A. Santoro,et al.  Activity, abundance and diversity of nitrifying archaea and bacteria in the central California Current. , 2010, Environmental microbiology.

[4]  Angelicque E. White,et al.  Unicellular Cyanobacterial Distributions Broaden the Oceanic N2 Fixation Domain , 2010, Science.

[5]  Jonathan P. Zehr,et al.  Metabolic streamlining in an open-ocean nitrogen-fixing cyanobacterium , 2010, Nature.

[6]  E. Delong,et al.  Abundances of crenarchaeal amoA genes and transcripts in the Pacific Ocean , 2010, Environmental microbiology.

[7]  F. Partensky,et al.  Prochlorococcus: advantages and limits of minimalism. , 2010, Annual review of marine science.

[8]  N. Mahowald,et al.  Impacts of atmospheric nutrient inputs on marine biogeochemistry , 2010 .

[9]  E. Virginia Armbrust,et al.  Iron and silicic acid effects on phytoplankton productivity, diversity, and chemical composition in the central equatorial Pacific Ocean , 2010 .

[10]  M. Kuypers,et al.  Water column anammox and denitrification in a temperate permanently stratified lake (Lake Rassnitzer, Germany). , 2009, Systematic and applied microbiology.

[11]  E. Achterberg,et al.  Large-scale distribution of Atlantic nitrogen fixation controlled by iron availability , 2009 .

[12]  V. Orphan,et al.  Deep-Sea Archaea Fix and Share Nitrogen in Methane-Consuming Microbial Consortia , 2009, Science.

[13]  D. Stahl,et al.  Ammonia oxidation kinetics determine niche separation of nitrifying Archaea and Bacteria , 2009, Nature.

[14]  J. Imhoff,et al.  Phylogenetic and functional marker genes to study ammonia-oxidizing microorganisms (AOM) in the environment , 2009, Applied Microbiology and Biotechnology.

[15]  B. Ward,et al.  Denitrification as the dominant nitrogen loss process in the Arabian Sea , 2009, Nature.

[16]  M. Kuypers,et al.  Anammox bacteria and the anaerobic oxidation of ammonium in the oxygen minimum zone off northern Chile , 2009 .

[17]  Adam C. Martiny,et al.  Widespread metabolic potential for nitrite and nitrate assimilation among Prochlorococcus ecotypes , 2009, Proceedings of the National Academy of Sciences.

[18]  C. Gobler,et al.  UREASE GENE SEQUENCES FROM ALGAE AND HETEROTROPHIC BACTERIA IN AXENIC AND NONAXENIC PHYTOPLANKTON CULTURES 1 , 2009, Journal of phycology.

[19]  Ricardo M Letelier,et al.  Physical forcing of nitrogen fixation and diazotroph community structure in the North Pacific subtropical gyre , 2009 .

[20]  M. Strous,et al.  Biochemistry and molecular biology of anammox bacteria , 2009, Critical reviews in biochemistry and molecular biology.

[21]  Edward F. DeLong,et al.  The microbial ocean from genomes to biomes , 2009, Nature.

[22]  I. Biegala,et al.  Diazotrophic unicellular cyanobacteria in the northwestern Mediterranean Sea: A seasonal cycle , 2009 .

[23]  S. Doney,et al.  Contributions of long-term research and time-series observations to marine ecology and biogeochemistry. , 2009, Annual review of marine science.

[24]  J. Zehr New twist on nitrogen cycling in oceanic oxygen minimum zones , 2009, Proceedings of the National Academy of Sciences.

[25]  R. Amann,et al.  Revising the nitrogen cycle in the Peruvian oxygen minimum zone , 2009, Proceedings of the National Academy of Sciences.

[26]  Lisa R. Moore,et al.  Phytoplankton in the ocean use non-phosphorus lipids in response to phosphorus scarcity , 2009, Nature.

[27]  R. Beinart,et al.  In situ transcriptomic analysis of the globally important keystone N2-fixing taxon Crocosphaera watsonii , 2009, The ISME Journal.

[28]  J. Gilbert,et al.  Nitrogen fixation in the western English Channel (NE Atlantic Ocean) , 2009 .

[29]  J. Dinasquet,et al.  Major gradients in putatively nitrifying and non-nitrifying Archaea in the deep North Atlantic , 2008, Nature.

[30]  P. Falkowski,et al.  Redox control of N:P ratios in aquatic ecosystems , 2008, Geobiology.

[31]  J. Montoya,et al.  Phylogenetic diversity of cyanobacterial narB genes from various marine habitats. , 2008, Environmental microbiology.

[32]  K. Fennel Widespread implementation of controlled upwelling in the North Pacific Subtropical Gyre would counteract diazotrophic N2 fixation , 2008 .

[33]  Ricardo M Letelier,et al.  Physical and ecological uncertainties in the widespread implementation of controlled upwelling in the North Pacific Subtropical Gyre , 2008 .

[34]  Jonathan P. Zehr,et al.  Globally Distributed Uncultivated Oceanic N2-Fixing Cyanobacteria Lack Oxygenic Photosystem II , 2008, Science.

[35]  Annika C. Mosier,et al.  Relative abundance and diversity of ammonia-oxidizing archaea and bacteria in the San Francisco Bay estuary. , 2008, Environmental microbiology.

[36]  Nicole L. Goebel,et al.  GROWTH AND CARBON CONTENT OF THREE DIFFERENT‐SIZED DIAZOTROPHIC CYANOBACTERIA OBSERVED IN THE SUBTROPICAL NORTH PACIFIC 1 , 2008, Journal of phycology.

[37]  G. Taylor,et al.  Vitamin B12 and cobalt cycling among diatoms and bacteria in Antarctic sea ice microbial communities , 2008 .

[38]  E. Carpenter,et al.  Amazon River enhances diazotrophy and carbon sequestration in the tropical North Atlantic Ocean , 2008, Proceedings of the National Academy of Sciences.

[39]  Ricardo M Letelier,et al.  Nitrogen fixation-enhanced carbon sequestration in low nitrate, low chlorophyll seascapes , 2008 .

[40]  J. Montoya,et al.  Recent advances in the biogeochemistry of nitrogen in the ocean , 2008 .

[41]  Deborah K. Steinberg,et al.  VERTIGO (VERtical Transport In the Global Ocean) : A study of particle sources and flux attenuation in the North Pacific , 2008 .

[42]  R. Beinart,et al.  Diversity and abundance of diazotrophic microorganisms in the South China Sea during intermonsoon , 2008, The ISME Journal.

[43]  F. Morel The co‐evolution of phytoplankton and trace element cycles in the oceans , 2008, Geobiology.

[44]  N. Jiao,et al.  Diversity and Abundance of Nitrate Assimilation Genes in the Northern South China Sea , 2008, Microbial Ecology.

[45]  J. Sprintall,et al.  Expanding Oxygen-Minimum Zones in the Tropical Oceans , 2008, Science.

[46]  M. D. de Jonge,et al.  Marine Polyphosphate: A Key Player in Geologic Phosphorus Sequestration , 2008, Science.

[47]  P. Raimbault,et al.  High abundance of diazotrophic picocyanobacteria (<3 µm) in a Southwest Pacific coral lagoon , 2008 .

[48]  James K. B. Bishop,et al.  The continental margin is a key source of iron to the HNLC North Pacific Ocean , 2008 .

[49]  J. Beman,et al.  Molecular and biogeochemical evidence for ammonia oxidation by marine Crenarchaeota in the Gulf of California , 2008, The ISME Journal.

[50]  Ricardo M Letelier,et al.  Nitrogen fixation in an anticyclonic eddy in the oligotrophic North Pacific Ocean , 2008, The ISME Journal.

[51]  J. LaRoche,et al.  Abundances and Distributions of the Dominant nifH Phylotypes in the Northern Atlantic Ocean , 2008, Applied and Environmental Microbiology.

[52]  B. Ward,et al.  Anaerobic Ammonium Oxidation (Anammox) in Chesapeake Bay Sediments , 2008, Microbial Ecology.

[53]  J. Galloway,et al.  An Earth-system perspective of the global nitrogen cycle , 2008, Nature.

[54]  B. V. Van Mooy,et al.  Assessing nutrient limitation of Prochlorococcus in the North Pacific subtropical gyre by using an RNA capture method , 2008 .

[55]  Ian Joint,et al.  Ammonium regeneration and nitrification rates in the oligotrophic Atlantic Ocean: Implications for new production estimates , 2007 .

[56]  E. Delong,et al.  Low genomic diversity in tropical oceanic N2-fixing cyanobacteria , 2007, Proceedings of the National Academy of Sciences.

[57]  Nicole L. Goebel,et al.  Modeled contributions of three types of diazotrophs to nitrogen fixation at Station ALOHA , 2007, The ISME Journal.

[58]  P. Raimbault,et al.  Nutrient limitation of primary productivity in the Southeast Pacific (BIOSOPE cruise) , 2007 .

[59]  C. Sakamoto,et al.  Nitrogen fixation by unicellular diazotrophic cyanobacteria in the temperate oligotrophic North Pacific Ocean , 2007 .

[60]  Scott C. Doney,et al.  Iron availability limits the ocean nitrogen inventory stabilizing feedbacks between marine denitrification and nitrogen fixation , 2007 .

[61]  D. Capone,et al.  What's New in the Nitrogen Cycle? , 2007 .

[62]  S. Giovannoni,et al.  New cultivation strategies bring more microbial plankton species into the laboratory , 2007 .

[63]  F. Chavez,et al.  Nitrification in the euphotic zone as evidenced by nitrate dual isotopic composition: Observations from Monterey Bay, California , 2007 .

[64]  D. Stahl,et al.  Functionally distinct communities of ammonia-oxidizing bacteria along an estuarine salinity gradient. , 2007, Environmental microbiology.

[65]  Nicholas R. Bates,et al.  Eddy/Wind Interactions Stimulate Extraordinary Mid-Ocean Plankton Blooms , 2007, Science.

[66]  M. Lomas,et al.  Prochlorococcus contributes to new production in the Sargasso Sea deep chlorophyll maximum , 2007 .

[67]  E. Delong,et al.  Quantitative distribution of presumptive archaeal and bacterial nitrifiers in Monterey Bay and the North Pacific Subtropical Gyre. , 2007, Environmental microbiology.

[68]  B. Quéguiner,et al.  Effect of natural iron fertilization on carbon sequestration in the Southern Ocean , 2007, Nature.

[69]  Sallie W. Chisholm,et al.  Emergent Biogeography of Microbial Communities in a Model Ocean , 2007, Science.

[70]  A. Halpern,et al.  The Sorcerer II Global Ocean Sampling Expedition: Northwest Atlantic through Eastern Tropical Pacific , 2007, PLoS biology.

[71]  E. Carpenter,et al.  Influence of the Amazon River plume on distributions of free‐living and symbiotic cyanobacteria in the western tropical north Atlantic Ocean , 2007 .

[72]  C. Deutsch,et al.  New developments in the marine nitrogen cycle. , 2007, Chemical reviews.

[73]  E. Boyle,et al.  Mesoscale Iron Enrichment Experiments 1993-2005: Synthesis and Future Directions , 2007, Science.

[74]  Nicolas Gruber,et al.  Spatial coupling of nitrogen inputs and losses in the ocean , 2007, Nature.

[75]  J. Baross,et al.  Nitrogen Fixation at 92°C by a Hydrothermal Vent Archaeon , 2006, Science.

[76]  D. Siegel,et al.  Spatial and temporal distribution of Trichodesmium blooms in the world's oceans , 2006 .

[77]  B. Jenkins,et al.  Cyanobacterial assimilatory nitrate reductase gene diversity in coastal and oligotrophic marine environments. , 2006, Environmental microbiology.

[78]  J. Montoya,et al.  Spatial variation in N 2 -fixation rate and diazotroph activity in the Tropical Atlantic , 2006 .

[79]  J. Zehr,et al.  Characterization of diatom-cyanobacteria symbioses on the basis of nifH, hetR and 16S rRNA sequences. , 2006, Environmental microbiology.

[80]  Ricardo M Letelier,et al.  Modeling carbohydrate ballasting by Trichodesmium spp. , 2006 .

[81]  Lily Shiue,et al.  Application of a nifH oligonucleotide microarray for profiling diversity of N2-fixing microorganisms in marine microbial mats. , 2006, Environmental microbiology.

[82]  R. Haselkorn,et al.  Genome sequence of Synechococcus CC9311: Insights into adaptation to a coastal environment , 2006, Proceedings of the National Academy of Sciences.

[83]  G. Rocap,et al.  Culture Isolation and Culture-Independent Clone Libraries Reveal New Marine Synechococcus Ecotypes with Distinctive Light and N Physiologies , 2006, Applied and Environmental Microbiology.

[84]  Marc Strous,et al.  Archaeal nitrification in the ocean , 2006, Proceedings of the National Academy of Sciences.

[85]  L. Codispoti An oceanic fixed nitrogen sink exceeding 400 Tg N a −1 vs the concept of homeostasis in the fixed-nitrogen inventory , 2006 .

[86]  Cabell S Davis,et al.  Transatlantic Abundance of the N2-Fixing Colonial Cyanobacterium Trichodesmium , 2006, Science.

[87]  Jorge L. Sarmiento,et al.  Ocean Biogeochemical Dynamics , 2006 .

[88]  Gabrielle Rocap,et al.  Sulfolipids dramatically decrease phosphorus demand by picocyanobacteria in oligotrophic marine environments. , 2006, Proceedings of the National Academy of Sciences of the United States of America.

[89]  J. Restrepo,et al.  A possible sequence of events for the generalized glacial‐interglacial cycle , 2006 .

[90]  Dmitrij Frishman,et al.  Deciphering the evolution and metabolism of an anammox bacterium from a community genome , 2006, Nature.

[91]  Sallie W. Chisholm,et al.  Niche Partitioning Among Prochlorococcus Ecotypes Along Ocean-Scale Environmental Gradients , 2006, Science.

[92]  G. Taylor,et al.  Regulation of phytoplankton dynamics by vitamin B12 , 2006 .

[93]  Wayne S Gardner,et al.  Nitrogen fixation and dissimilatory nitrate reduction to ammonium (DNRA) support nitrogen dynamics in Texas estuaries , 2006 .

[94]  K. Lindsay,et al.  Nitrogen fixation amplifies the ocean biogeochemical response to decadal timescale variations in mineral dust deposition , 2006 .

[95]  Hans-Peter Klenk,et al.  Novel genes for nitrite reductase and Amo-related proteins indicate a role of uncultivated mesophilic crenarchaeota in nitrogen cycling. , 2005, Environmental microbiology.

[96]  J. Sarmiento,et al.  Empirical and mechanistic models for the particle export ratio , 2005 .

[97]  J. Beman,et al.  Ubiquity and diversity of ammonia-oxidizing archaea in water columns and sediments of the ocean. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[98]  M. Könneke,et al.  Isolation of an autotrophic ammonia-oxidizing marine archaeon , 2005, Nature.

[99]  S. Giovannoni,et al.  Molecular diversity and ecology of microbial plankton , 2005, Nature.

[100]  K. Arrigo Marine microorganisms and global nutrient cycles , 2005, Nature.

[101]  E. Carpenter,et al.  Picoplankton community structure within and outside a Trichodesmium bloom in the southwestern Pacific Ocean , 2005 .

[102]  C. M. Short,et al.  Temporal Patterns of Nitrogenase Gene (nifH) Expression in the Oligotrophic North Pacific Ocean , 2005, Applied and Environmental Microbiology.

[103]  M. Noordewier,et al.  Genome Streamlining in a Cosmopolitan Oceanic Bacterium , 2005, Science.

[104]  A. Allen,et al.  Influence of nitrate availability on the distribution and abundance of heterotrophic bacterial nitrate assimilation genes in the Barents Sea during summer , 2005 .

[105]  E. Carpenter,et al.  Nitrogen fixation by Trichodesmium spp.: An important source of new nitrogen to the tropical and subtropical North Atlantic Ocean , 2005 .

[106]  C. Mahaffey The conundrum of marine N2 fixation , 2005 .

[107]  B. Ward Temporal variability in nitrification rates and related biogeochemical factors in Monterey Bay, California, USA , 2005 .

[108]  R. Amann,et al.  Massive nitrogen loss from the Benguela upwelling system through anaerobic ammonium oxidation. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[109]  M. Wagner,et al.  Biomarkers for In Situ Detection of Anaerobic Ammonium-Oxidizing (Anammox) Bacteria , 2005, Applied and Environmental Microbiology.

[110]  N. Mahowald,et al.  Global Iron Connections Between Desert Dust, Ocean Biogeochemistry, and Climate , 2005, Science.

[111]  E. Boyle,et al.  Iron, manganese, and lead at Hawaii Ocean Time-series station ALOHA: Temporal variability and an intermediate water hydrothermal plume , 2005 .

[112]  D. Karl,et al.  Vertical distributions of nitrogen-fixing phylotypes at Stn ALOHA in the oligotrophic North Pacific Ocean , 2005 .

[113]  D. Capone,et al.  Dinitrogen fixation and release of ammonium and dissolved organic nitrogen by Trichodesmium IMS101 , 2004 .

[114]  S. Doney,et al.  From genes to ecosystems: the ocean's new frontier , 2004 .

[115]  G. Asner,et al.  Nitrogen Cycles: Past, Present, and Future , 2004 .

[116]  Matthew M. Mills,et al.  Iron and phosphorus co-limit nitrogen fixation in the eastern tropical North Atlantic , 2004, Nature.

[117]  S. Levin,et al.  Optimal nitrogen-to-phosphorus stoichiometry of phytoplankton , 2004, Nature.

[118]  O. White,et al.  Environmental Genome Shotgun Sequencing of the Sargasso Sea , 2004, Science.

[119]  Bess B. Ward,et al.  Development and Testing of a DNA Macroarray To Assess Nitrogenase (nifH) Gene Diversity , 2004, Applied and Environmental Microbiology.

[120]  G. Steward,et al.  Fingerprinting Diazotroph Communities in the Chesapeake Bay by Using a DNA Macroarray , 2004, Applied and Environmental Microbiology.

[121]  F. Cipriano,et al.  N2 Fixation by Unicellular Bacterioplankton from the Atlantic and Pacific Oceans: Phylogeny and In Situ Rates , 2004, Applied and Environmental Microbiology.

[122]  M. Church,et al.  Nitrogenase genes in PCR and RT-PCR reagents: implications for studies of diversity of functional genes. , 2003, BioTechniques.

[123]  Manesh Shah,et al.  Genome divergence in two Prochlorococcus ecotypes reflects oceanic niche differentiation , 2003, Nature.

[124]  Michael Y. Galperin,et al.  Genome sequence of the cyanobacterium Prochlorococcus marinus SS120, a nearly minimal oxyphototrophic genome , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[125]  F. Morel,et al.  The Biogeochemical Cycles of Trace Metals in the Oceans , 2003, Science.

[126]  D. Karl,et al.  Bioavailability of dissolved organic phosphorus in the euphotic zone at Station ALOHA, North Pacific Subtropical Gyre , 2003 .

[127]  J. G. Kuenen,et al.  Anaerobic ammonium oxidation by anammox bacteria in the Black Sea , 2003, Nature.

[128]  D. Harbour,et al.  Large-scale latitudinal distribution of Trichodesmium spp. in the Atlantic Ocean , 2003 .

[129]  E. Carpenter,et al.  A REVISED ESTIMATE OF THE IRON USE EFFICIENCY OF NITROGEN FIXATION, WITH SPECIAL REFERENCE TO THE MARINE CYANOBACTERIUM TRICHODESMIUM SPP. (CYANOPHYTA) 1 , 2003 .

[130]  Lars-Anders Hansson,et al.  Cell‐specific detection of phosphorus stress in Trichodesmium from the Western North Atlantic , 2002 .

[131]  D. Karl,et al.  Nutrient dynamics in the deep blue sea. , 2002, Trends in microbiology.

[132]  B. Thamdrup,et al.  Production of N2 through Anaerobic Ammonium Oxidation Coupled to Nitrate Reduction in Marine Sediments , 2002, Applied and Environmental Microbiology.

[133]  H. Ducklow,et al.  Widespread Distribution in Polar Oceans of a 16S rRNA Gene Sequence with Affinity to Nitrosospira-Like Ammonia-Oxidizing Bacteria , 2002, Applied and Environmental Microbiology.

[134]  R. Geider,et al.  Redfield revisited: variability of C:N:P in marine microalgae and its biochemical basis , 2002 .

[135]  M. Frischer,et al.  Diversity and Detection of Nitrate Assimilation Genes in Marine Bacteria , 2001, Applied and Environmental Microbiology.

[136]  Andrew Hansen,et al.  Unicellular cyanobacteria fix N2 in the subtropical North Pacific Ocean , 2001, Nature.

[137]  C. Gobler,et al.  Phosphorus limitation of nitrogen fixation by Trichodesmium in the central Atlantic Ocean , 2001, Nature.

[138]  J. Beardall,et al.  Approaches for determining phytoplankton nutrient limitation , 2001, Aquatic Sciences.

[139]  William F. Fagan,et al.  Biological stoichiometry from genes to ecosystems. , 2000 .

[140]  E. Boyle,et al.  Phosphate depletion in the western North Atlantic Ocean. , 2000, Science.

[141]  Edward J Carpenter,et al.  INTRACELLULAR CYANOBACTERIAL SYMBIONTS IN THE MARINE DIATOM CLIMACODIUM FRAUENFELDIANUM (BACILLARIOPHYCEAE) , 2000, Journal of phycology.

[142]  S. Doney,et al.  Iron supply and demand in the upper ocean , 2000 .

[143]  Toby Tyrrell,et al.  The relative influences of nitrogen and phosphorus on oceanic primary production , 1999, Nature.

[144]  D. Karl,et al.  Diatom fluxes to the deep sea in the oligotrophic North Pacific gyre at Station ALOHA , 1999 .

[145]  D. Vaulot,et al.  Phycoerythrins in the southern tropical and equatorial Pacific Ocean: Evidence for new cyanobacterial types , 1999 .

[146]  M. Behrenfeld,et al.  Widespread iron limitation of phytoplankton in the south pacific ocean , 1999, Science.

[147]  Jonathan P. Zehr,et al.  New Nitrogen-Fixing Microorganisms Detected in Oligotrophic Oceans by Amplification of Nitrogenase (nifH) Genes , 1998, Applied and Environmental Microbiology.

[148]  Lisa R. Moore,et al.  Physiology and molecular phylogeny of coexisting Prochlorococcus ecotypes , 1998, Nature.

[149]  G. Likens,et al.  Technical Report: Human Alteration of the Global Nitrogen Cycle: Sources and Consequences , 1997 .

[150]  J. G. Kuenen,et al.  Metabolic pathway of anaerobic ammonium oxidation on the basis of 15N studies in a fluidized bed reactor. , 1997, Microbiology.

[151]  Nicolas Gruber,et al.  Global patterns of marine nitrogen fixation and denitrification , 1997 .

[152]  J. Downing Marine nitrogen: Phosphorus stoichiometry and the global N:P cycle , 1997 .

[153]  Paul G. Falkowski,et al.  Evolution of the nitrogen cycle and its influence on the biological sequestration of CO2 in the ocean , 1997, Nature.

[154]  D. Karl,et al.  Nitrification in the euphotic zone as a source for nitrite, nitrate, and nitrous oxide at Station ALOHA , 1996 .

[155]  B. Ward Nitrification and denitrification: Probing the nitrogen cycle in aquatic environments , 1996, Microbial Ecology.

[156]  Lisa M. Graziano,et al.  Nitrogen limitation of North Atlantic phytoplankton: analysis of physiological condition in nutrient enrichment experiments , 1996 .

[157]  B. Bergman,et al.  Intracellular cyanobiont Richelia intracellularis: ultrastructure and immuno-localisation of phycoerythrin, nitrogenase, Rubisco and glutamine synthetase , 1995 .

[158]  J. G. Kuenen,et al.  Anaerobic ammonium oxidation discovered in a denitrifying fluidized bed reactor , 1995 .

[159]  M. Altabet,et al.  Nitrogen transport by vertically migrating diatom mats in the North Pacific Ocean , 1993, Nature.

[160]  E. Delong Archaea in coastal marine environments. , 1992, Proceedings of the National Academy of Sciences of the United States of America.

[161]  A. Davis Novel major archaebacterial group from marine plankton , 1992, Nature.

[162]  R. Eppley,et al.  Biological nitrogen cycling in the nitracline , 1989 .

[163]  R. Hecky,et al.  Nutrient limitation of phytoplankton in freshwater and marine environments: A review of recent evidence on the effects of enrichment1 , 1988 .

[164]  Sallie W. Chisholm,et al.  A novel free-living prochlorophyte abundant in the oceanic euphotic zone , 1988, Nature.

[165]  Cécile Dupouy,et al.  Satellite detected cyanobacteria bloom in the southwestern tropical Pacific Implication for oceanic nitrogen fixation , 1988 .

[166]  B. Ward,et al.  Marine Ammonia- and Nitrite-Oxidizing Bacteria: Serological Diversity Determined by Immunofluorescence in Culture and in the Environment , 1985, Applied and environmental microbiology.

[167]  A. Devol,et al.  Nitrogen in the Marine Environment , 1985 .

[168]  S. V. Smith Phosphorus versus nitrogen limitation in the marine environment1 , 1984 .

[169]  B. Peterson,et al.  Particulate organic matter flux and planktonic new production in the deep ocean , 1979, Nature.

[170]  T. Saino,et al.  Diel variation in nitrogen fixation by a marine blue-green alga, Trichodesmium thiebautii , 1978 .

[171]  O. Holm‐Hansen,et al.  Nitrogen fixation in the North Pacific Ocean , 1974 .

[172]  J. Ryther,et al.  Nitrogen, Phosphorus, and Eutrophication in the Coastal Marine Environment , 1971, Science.

[173]  W. Thomas ON NITROGEN DEFICIENCY IN TROPICAL PACIFIC OCEANIC PHYTOPLANKTON: PHOTOSYNTHETIC PARAMETERS IN POOR AND RICH WATER1 , 1970 .

[174]  J. Goering,et al.  UPTAKE OF NEW AND REGENERATED FORMS OF NITROGEN IN PRIMARY PRODUCTIVITY1 , 1967 .

[175]  J. Ryther,et al.  Nitrogen fixation in the Sargasso Sea , 1961 .

[176]  R. Anadón,et al.  Trichodesmium spp. population structure in the eastern North Atlantic subtropical gyre , 2010 .

[177]  M. Strous,et al.  Presence and activity of anaerobic ammonium-oxidizing bacteria at deep-sea hydrothermal vents , 2009, The ISME Journal.

[178]  M. Follows,et al.  On the interannual variability of nitrogen fixation in the subtropical gyres , 2009 .

[179]  M. McCarthy,et al.  Analytical Methods for the Study of Nitrogen , 2008 .

[180]  R. Foster,et al.  Nitrogen-Fixing and Nitrifying Symbioses in the Marine Environment , 2008 .

[181]  L. Riemann,et al.  Isolation and gene quantification of heterotrophic N2-fixing bacterioplankton in the Baltic Sea. , 2007, Environmental microbiology.

[182]  M. Kuypers,et al.  Anaerobic ammonium oxidation in the marine environment , 2006 .

[183]  Nicholas R. Bates,et al.  Excess nitrate and nitrogen fixation in the North Atlantic Ocean , 2004 .

[184]  Ricardo M Letelier,et al.  Ecological nitrogen-to-phosphorus stoichiometry at station ALOHA , 2001 .

[185]  R. Bidigare,et al.  Long-term changes in plankton community structure and productivity in the North Pacific Subtropical Gyre: The domain shift hypothesis , 2001 .

[186]  A. Rees,et al.  Nutrient distributions in an anticyclonic eddy in the northeast Atlantic Ocean, with reference to nanomolar ammonium concentrations , 2001 .

[187]  E. Carpenter,et al.  Detecting Trichodesmium blooms in SeaWiFS imagery , 2001 .

[188]  D. Karl,et al.  Microbial ecology at sea: Sampling, subsampling and incubation considerations , 2001 .

[189]  M. Pahlow,et al.  Temporal trends in deep ocean Redfield ratios. , 2000, Science.

[190]  J. Sarmiento,et al.  Inputs, losses and transformations of nitrogen and phosphorus in the pelagic North Atlantic Ocean , 1996 .

[191]  P. Falkowski,et al.  Iron limitation of phytoplankton photosynthesis in the equatorial Pacific Ocean , 1994, Nature.

[192]  T. A. Villareal,et al.  Marine Nitrogen-Fixing Diatom-Cyanobacteria Symbioses , 1992 .

[193]  Ricardo M Letelier,et al.  Tricho-desmium blooms and new nitrogen in the North Pacific gyre , 1992 .

[194]  Tracy A. Villareal,et al.  Nitrogen-fixation by the cyanobacterial symbiont of the diatom genus Hemiaulus , 1991 .

[195]  Robert W. Howarth,et al.  Nitrogen limitation on land and in the sea: How can it occur? , 1991 .

[196]  S. Fitzwater,et al.  Iron deficiency limits phytoplankton growth in the north-east Pacific subarctic , 1988, Nature.

[197]  R. Howarth Nutrient Limitation of Net Primary Production in Marine Ecosystems , 1988 .

[198]  Tracy A. Villareal,et al.  Evaluation of nitrogen fixation in the diatom genus Rhizosolenia Ehr. in the absence of its cyanobacterial symbiont Richelia intracellularis Schmidt , 1987 .

[199]  E. Carpenter Chapter 3 – NITROGEN FIXATION BY MARINE OSCILLATORIA (TRICHODESMIUM) IN THE WORLD's OCEANS , 1983 .

[200]  A. C. Redfield The biological control of chemical factors in the environment. , 1960, Science progress.