Articles | Volume 17, issue 8
https://doi.org/10.5194/bg-17-2263-2020
© Author(s) 2020. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
https://doi.org/10.5194/bg-17-2263-2020
© Author(s) 2020. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Regulation of nitrous oxide production in low-oxygen waters off the coast of Peru
Department of Geoscience, Princeton University, Princeton, Guyot Hall, Princeton, NJ 08544, USA
Helmholtz Centre for Ocean Research Kiel, Düsternbrooker Weg 20, 24105 Kiel, Germany
current address: Department of Environmental Science, University of Basel,
Bernoullistrasse 30, 4056 Basel, Switzerland
Hermann W. Bange
Helmholtz Centre for Ocean Research Kiel, Düsternbrooker Weg 20, 24105 Kiel, Germany
Eric P. Achterberg
Helmholtz Centre for Ocean Research Kiel, Wischhofstr. 1–3, 24149 Kiel, Germany
Amal Jayakumar
Department of Geoscience, Princeton University, Princeton, Guyot Hall, Princeton, NJ 08544, USA
Carolin R. Löscher
Department of Biology, Nordcee, Danish Institute for Advanced Study, University of Southern Denmark, Odense, Denmark
Damian L. Arévalo-Martínez
Helmholtz Centre for Ocean Research Kiel, Düsternbrooker Weg 20, 24105 Kiel, Germany
Elizabeth León-Palmero
Departamento de Ecología, Facultad de Ciencias, Universidad de Granada, 18071, Granada, Spain
Mingshuang Sun
Helmholtz Centre for Ocean Research Kiel, Düsternbrooker Weg 20, 24105 Kiel, Germany
Xin Sun
Department of Geoscience, Princeton University, Princeton, Guyot Hall, Princeton, NJ 08544, USA
Ruifang C. Xie
Helmholtz Centre for Ocean Research Kiel, Wischhofstr. 1–3, 24149 Kiel, Germany
Sergey Oleynik
Department of Geoscience, Princeton University, Princeton, Guyot Hall, Princeton, NJ 08544, USA
Bess B. Ward
Department of Geoscience, Princeton University, Princeton, Guyot Hall, Princeton, NJ 08544, USA
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Jakob Rønning, Zarah J. Kofoed, Mats Jacobsen, and Carolin R. Löscher
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Zhibo Shao, Yangchun Xu, Hua Wang, Weicheng Luo, Lice Wang, Yuhong Huang, Nona Sheila R. Agawin, Ayaz Ahmed, Mar Benavides, Mikkel Bentzon-Tilia, Ilana Berman-Frank, Hugo Berthelot, Isabelle C. Biegala, Mariana B. Bif, Antonio Bode, Sophie Bonnet, Deborah A. Bronk, Mark V. Brown, Lisa Campbell, Douglas G. Capone, Edward J. Carpenter, Nicolas Cassar, Bonnie X. Chang, Dreux Chappell, Yuh-ling Lee Chen, Matthew J. Church, Francisco M. Cornejo-Castillo, Amália Maria Sacilotto Detoni, Scott C. Doney, Cecile Dupouy, Marta Estrada, Camila Fernandez, Bieito Fernández-Castro, Debany Fonseca-Batista, Rachel A. Foster, Ken Furuya, Nicole Garcia, Kanji Goto, Jesús Gago, Mary R. Gradoville, M. Robert Hamersley, Britt A. Henke, Cora Hörstmann, Amal Jayakumar, Zhibing Jiang, Shuh-Ji Kao, David M. Karl, Leila R. Kittu, Angela N. Knapp, Sanjeev Kumar, Julie LaRoche, Hongbin Liu, Jiaxing Liu, Caroline Lory, Carolin R. Löscher, Emilio Marañón, Lauren F. Messer, Matthew M. Mills, Wiebke Mohr, Pia H. Moisander, Claire Mahaffey, Robert Moore, Beatriz Mouriño-Carballido, Margaret R. Mulholland, Shin-ichiro Nakaoka, Joseph A. Needoba, Eric J. Raes, Eyal Rahav, Teodoro Ramírez-Cárdenas, Christian Furbo Reeder, Lasse Riemann, Virginie Riou, Julie C. Robidart, Vedula V. S. S. Sarma, Takuya Sato, Himanshu Saxena, Corday Selden, Justin R. Seymour, Dalin Shi, Takuhei Shiozaki, Arvind Singh, Rachel E. Sipler, Jun Sun, Koji Suzuki, Kazutaka Takahashi, Yehui Tan, Weiyi Tang, Jean-Éric Tremblay, Kendra Turk-Kubo, Zuozhu Wen, Angelicque E. White, Samuel T. Wilson, Takashi Yoshida, Jonathan P. Zehr, Run Zhang, Yao Zhang, and Ya-Wei Luo
Earth Syst. Sci. Data, 15, 3673–3709, https://doi.org/10.5194/essd-15-3673-2023, https://doi.org/10.5194/essd-15-3673-2023, 2023
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Nitrous oxide (N2O) is an important greenhouse gas. However, N2O emissions from estuaries underlie significant uncertainties due to limited data availability and high spatiotemporal variability. We found the Elbe Estuary (Germany) to be a year-round source of N2O, with the highest emissions in winter along with high nitrogen loads. However, in spring and summer, N2O emissions did not decrease alongside lower nitrogen loads because organic matter fueled in situ N2O production along the estuary.
John C. Tracey, Andrew R. Babbin, Elizabeth Wallace, Xin Sun, Katherine L. DuRussel, Claudia Frey, Donald E. Martocello III, Tyler Tamasi, Sergey Oleynik, and Bess B. Ward
Biogeosciences, 20, 2499–2523, https://doi.org/10.5194/bg-20-2499-2023, https://doi.org/10.5194/bg-20-2499-2023, 2023
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Nitrogen (N) is essential for life; thus, its availability plays a key role in determining marine productivity. Using incubations of seawater spiked with a rare form of N measurable on a mass spectrometer, we quantified microbial pathways that determine marine N availability. The results show that pathways that recycle N have higher rates than those that result in its loss from biomass and present new evidence for anaerobic nitrite oxidation, a process long thought to be strictly aerobic.
Guanlin Li, Damian L. Arévalo-Martínez, Riel Carlo O. Ingeniero, and Hermann W. Bange
EGUsphere, https://doi.org/10.5194/egusphere-2023-771, https://doi.org/10.5194/egusphere-2023-771, 2023
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Dissolved carbon monoxide (CO) surface concentrations were first measured at 14 stations in the Ria Formosa Lagoon system in May 2021. Ria Formosa was a source of atmospheric CO. Microbial consumption accounted for 83 % of the CO production. The results of a 48-hour irradiation experiment with aquaculture effluent water indicated that aquaculture facilities in the Ria Formosa Lagoon seem to be a negligible source of atmospheric CO.
Kristian Spilling, Jonna Piiparinen, Eric P. Achterberg, Javier Arístegui, Lennart T. Bach, Maria T. Camarena-Gómez, Elisabeth von der Esch, Martin A. Fischer, Markel Gómez-Letona, Nauzet Hernández-Hernández, Judith Meyer, Ruth A. Schmitz, and Ulf Riebesell
Biogeosciences, 20, 1605–1619, https://doi.org/10.5194/bg-20-1605-2023, https://doi.org/10.5194/bg-20-1605-2023, 2023
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We carried out an enclosure experiment using surface water off Peru with different additions of oxygen minimum zone water. In this paper, we report on enzyme activity and provide data on the decomposition of organic matter. We found very high activity with respect to an enzyme breaking down protein, suggesting that this is important for nutrient recycling both at present and in the future ocean.
Hanna I. Campen, Damian L. Arévalo-Martínez, and Hermann W. Bange
Biogeosciences, 20, 1371–1379, https://doi.org/10.5194/bg-20-1371-2023, https://doi.org/10.5194/bg-20-1371-2023, 2023
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Carbon monoxide (CO) is a climate-relevant trace gas emitted from the ocean. However, oceanic CO cycling is understudied. Results from incubation experiments conducted in the Fram Strait (Arctic Ocean) indicated that (i) pH did not affect CO cycling and (ii) enhanced CO production and consumption were positively correlated with coloured dissolved organic matter and nitrate concentrations. This suggests microbial CO uptake to be the driving factor for CO cycling in the Arctic Ocean.
Damian L. Arévalo-Martínez, Amir Haroon, Hermann W. Bange, Ercan Erkul, Marion Jegen, Nils Moosdorf, Jens Schneider von Deimling, Christian Berndt, Michael Ernst Böttcher, Jasper Hoffmann, Volker Liebetrau, Ulf Mallast, Gudrun Massmann, Aaron Micallef, Holly A. Michael, Hendrik Paasche, Wolfgang Rabbel, Isaac Santos, Jan Scholten, Katrin Schwalenberg, Beata Szymczycha, Ariel T. Thomas, Joonas J. Virtasalo, Hannelore Waska, and Bradley A. Weymer
Biogeosciences, 20, 647–662, https://doi.org/10.5194/bg-20-647-2023, https://doi.org/10.5194/bg-20-647-2023, 2023
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Groundwater flows at the land–ocean transition and the extent of freshened groundwater below the seafloor are increasingly relevant in marine sciences, both because they are a highly uncertain term of biogeochemical budgets and due to the emerging interest in the latter as a resource. Here, we discuss our perspectives on future research directions to better understand land–ocean connectivity through groundwater and its potential responses to natural and human-induced environmental changes.
Sonja Gindorf, Hermann W. Bange, Dennis Booge, and Annette Kock
Biogeosciences, 19, 4993–5006, https://doi.org/10.5194/bg-19-4993-2022, https://doi.org/10.5194/bg-19-4993-2022, 2022
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Methane is a climate-relevant greenhouse gas which is emitted to the atmosphere from coastal areas such as the Baltic Sea. We measured the methane concentration in the water column of the western Kiel Bight. Methane concentrations were higher in September than in June. We found no relationship between the 2018 European heatwave and methane concentrations. Our results show that the methane distribution in the water column is strongly affected by temporal and spatial variabilities.
Mhlangabezi Mdutyana, Tanya Marshall, Xin Sun, Jessica M. Burger, Sandy J. Thomalla, Bess B. Ward, and Sarah E. Fawcett
Biogeosciences, 19, 3425–3444, https://doi.org/10.5194/bg-19-3425-2022, https://doi.org/10.5194/bg-19-3425-2022, 2022
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Nitrite-oxidizing bacteria in the winter Southern Ocean show a high affinity for nitrite but require a minimum (i.e., "threshold") concentration before they increase their rates of nitrite oxidation significantly. The classic Michaelis–Menten model thus cannot be used to derive the kinetic parameters, so a modified equation was employed that also yields the threshold nitrite concentration. Dissolved iron availability may play an important role in limiting nitrite oxidation.
Christian Furbo Reeder, Ina Stoltenberg, Jamileh Javidpour, and Carolin Regina Löscher
Ocean Sci., 18, 401–417, https://doi.org/10.5194/os-18-401-2022, https://doi.org/10.5194/os-18-401-2022, 2022
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The Baltic Sea is predicted to freshen in the future. To explore the effect of decreasing salinity on N2 fixers, we followed the natural salinity gradient in the Baltic Sea from the Kiel Fjord to the Gotland Basin and identified an N2 fixer community dominated by Nodularia and UCYN-A. A salinity threshold was identified at a salinity of 10, with Nodularia dominating at low and UCYN-A dominating at higher salinity, suggesting a future expansion of Nodularia N2 fixers and a retraction of UCYN-A.
Yanan Zhao, Dennis Booge, Christa A. Marandino, Cathleen Schlundt, Astrid Bracher, Elliot L. Atlas, Jonathan Williams, and Hermann W. Bange
Biogeosciences, 19, 701–714, https://doi.org/10.5194/bg-19-701-2022, https://doi.org/10.5194/bg-19-701-2022, 2022
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We present here, for the first time, simultaneously measured dimethylsulfide (DMS) seawater concentrations and DMS atmospheric mole fractions from the Peruvian upwelling region during two cruises in December 2012 and October 2015. Our results indicate low oceanic DMS concentrations and atmospheric DMS molar fractions in surface waters and the atmosphere, respectively. In addition, the Peruvian upwelling region was identified as an insignificant source of DMS emissions during both periods.
Shao-Min Chen, Ulf Riebesell, Kai G. Schulz, Elisabeth von der Esch, Eric P. Achterberg, and Lennart T. Bach
Biogeosciences, 19, 295–312, https://doi.org/10.5194/bg-19-295-2022, https://doi.org/10.5194/bg-19-295-2022, 2022
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Oxygen minimum zones in the ocean are characterized by enhanced carbon dioxide (CO2) levels and are being further acidified by increasing anthropogenic atmospheric CO2. Here we report CO2 system measurements in a mesocosm study offshore Peru during a rare coastal El Niño event to investigate how CO2 dynamics may respond to ongoing ocean deoxygenation. Our observations show that nitrogen limitation, productivity, and plankton community shift play an important role in driving the CO2 dynamics.
Wangwang Ye, Hermann W. Bange, Damian L. Arévalo-Martínez, Hailun He, Yuhong Li, Jianwen Wen, Jiexia Zhang, Jian Liu, Man Wu, and Liyang Zhan
Biogeosciences Discuss., https://doi.org/10.5194/bg-2021-334, https://doi.org/10.5194/bg-2021-334, 2022
Manuscript not accepted for further review
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CH4 is the second important greenhouse gas after CO2. We show that CH4 consumption and sea-ice melting influence the CH4 distribution in the Ross Sea (Southern Ocean), causing undersaturation and net uptake of CH4 during summertime. This study confirms the capability of surface water in the high-latitude Southern Ocean regions to take up atmospheric CH4 which, in turn, will help to improve predictions of how CH4 release/uptake from the ocean will develop when sea-ice retreats in the future.
Carolin R. Löscher
Biogeosciences, 18, 4953–4963, https://doi.org/10.5194/bg-18-4953-2021, https://doi.org/10.5194/bg-18-4953-2021, 2021
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The Bay of Bengal (BoB) is classically seen as an ocean region with low primary production, which has been predicted to decrease even further. Here, the importance of such a trend is used to explore what could happen to the BoB's low-oxygen core waters if primary production decreases. Lower biological production leads to less oxygen loss in deeper waters by respiration; thus it could be that oxygen will not further decrease and the BoB will not become anoxic, different to other low-oxygen areas.
Kai G. Schulz, Eric P. Achterberg, Javier Arístegui, Lennart T. Bach, Isabel Baños, Tim Boxhammer, Dirk Erler, Maricarmen Igarza, Verena Kalter, Andrea Ludwig, Carolin Löscher, Jana Meyer, Judith Meyer, Fabrizio Minutolo, Elisabeth von der Esch, Bess B. Ward, and Ulf Riebesell
Biogeosciences, 18, 4305–4320, https://doi.org/10.5194/bg-18-4305-2021, https://doi.org/10.5194/bg-18-4305-2021, 2021
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Upwelling of nutrient-rich deep waters to the surface make eastern boundary upwelling systems hot spots of marine productivity. This leads to subsurface oxygen depletion and the transformation of bioavailable nitrogen into inert N2. Here we quantify nitrogen loss processes following a simulated deep water upwelling. Denitrification was the dominant process, and budget calculations suggest that a significant portion of nitrogen that could be exported to depth is already lost in the surface ocean.
Neil J. Wyatt, Angela Milne, Eric P. Achterberg, Thomas J. Browning, Heather A. Bouman, E. Malcolm S. Woodward, and Maeve C. Lohan
Biogeosciences, 18, 4265–4280, https://doi.org/10.5194/bg-18-4265-2021, https://doi.org/10.5194/bg-18-4265-2021, 2021
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Using data collected during two expeditions to the South Atlantic Ocean, we investigated how the interaction between external sources and biological activity influenced the availability of the trace metals zinc and cobalt. This is important as both metals play essential roles in the metabolism and growth of phytoplankton and thus influence primary productivity of the oceans. We found seasonal changes in both processes that helped explain upper-ocean trace metal cycling.
Siqi Wu, Moge Du, Xianhui Sean Wan, Corday Selden, Mar Benavides, Sophie Bonnet, Robert Hamersley, Carolin R. Löscher, Margaret R. Mulholland, Xiuli Yan, and Shuh-Ji Kao
Biogeosciences Discuss., https://doi.org/10.5194/bg-2021-104, https://doi.org/10.5194/bg-2021-104, 2021
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Nitrogen (N2) fixation is one of the most important nutrient sources to the ocean. Here, we report N2 fixation in the deep, dark ocean in the South China Sea via a highly sensitive new method and elaborate controls, showing the overlooked importance of N2 fixation in the deep ocean. By global data compilation, we also provide an easy measured basic parameter to estimate deep N2 fixation. Our study may help to expand the area limit of N2 fixation studies and better constrain global N2 fixation.
Maximiliano J. Vergara-Jara, Mark J. Hopwood, Thomas J. Browning, Insa Rapp, Rodrigo Torres, Brian Reid, Eric P. Achterberg, and José Luis Iriarte
Ocean Sci., 17, 561–578, https://doi.org/10.5194/os-17-561-2021, https://doi.org/10.5194/os-17-561-2021, 2021
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Ash from the Calbuco 2015 eruption spread across northern Patagonia, the SE Pacific and the SW Atlantic. In the Pacific, a phytoplankton bloom corresponded closely to the volcanic ash plume, suggesting that ash fertilized this region of the ocean. No such fertilization was found in the Atlantic where nutrients plausibly supplied by ash were likely already in excess of phytoplankton demand. In Patagonia, the May bloom was more intense than usual, but the mechanistic link to ash was less clear.
Gerd Krahmann, Damian L. Arévalo-Martínez, Andrew W. Dale, Marcus Dengler, Anja Engel, Nicolaas Glock, Patricia Grasse, Johannes Hahn, Helena Hauss, Mark Hopwood, Rainer Kiko, Alexandra Loginova, Carolin R. Löscher, Marie Maßmig, Alexandra-Sophie Roy, Renato Salvatteci, Stefan Sommer, Toste Tanhua, and Hela Mehrtens
Earth Syst. Sci. Data Discuss., https://doi.org/10.5194/essd-2020-308, https://doi.org/10.5194/essd-2020-308, 2021
Preprint withdrawn
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The project "Climate-Biogeochemistry Interactions in the Tropical Ocean" (SFB 754) was a multidisciplinary research project active from 2008 to 2019 aimed at a better understanding of the coupling between the tropical climate and ocean circulation and the ocean's oxygen and nutrient balance. On 34 research cruises, mainly in the Southeast Tropical Pacific and the Northeast Tropical Atlantic, 1071 physical, chemical and biological data sets were collected.
Yanan Zhao, Cathleen Schlundt, Dennis Booge, and Hermann W. Bange
Biogeosciences, 18, 2161–2179, https://doi.org/10.5194/bg-18-2161-2021, https://doi.org/10.5194/bg-18-2161-2021, 2021
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We present a unique and comprehensive time-series study of biogenic sulfur compounds in the southwestern Baltic Sea, from 2009 to 2018. Dimethyl sulfide is one of the key players regulating global climate change, as well as dimethylsulfoniopropionate and dimethyl sulfoxide. Their decadal trends did not follow increasing temperature but followed some algae group abundances at the Boknis Eck Time Series Station.
Yu-Te Hsieh, Walter Geibert, E. Malcolm S. Woodward, Neil J. Wyatt, Maeve C. Lohan, Eric P. Achterberg, and Gideon M. Henderson
Biogeosciences, 18, 1645–1671, https://doi.org/10.5194/bg-18-1645-2021, https://doi.org/10.5194/bg-18-1645-2021, 2021
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The South Atlantic near 40° S is one of the high-productivity and most dynamic nutrient regions in the oceans, but the sources and fluxes of trace elements (TEs) to this region remain unclear. This study investigates seawater Ra-228 and provides important constraints on ocean mixing and dissolved TE fluxes to this region. Vertical mixing is a more important source than aeolian or shelf inputs in this region, but particulate or winter deep-mixing inputs may be required to balance the TE budgets.
Cited articles
Anderson, J. H.: The metabolism of hydroxylamine to nitrite by Nitrosomonas,
Biochem. J., 91, 8–17, https://doi.org/10.1042/bj0910008, 1964.
Arévalo-Martínez, D. L., Kock, A., Löscher, C. R., Schmitz, R.
A., Bange, H. W., Arevalo-Martínez, D., Kock, A., Löscher, C. R.,
Schmitz, R. A., and Bange, H. W.: Massive nitrous oxide emissions from the
tropical South Pacific Ocean, Nat. Geosci., 8, 530–533,
https://doi.org/10.1038/NGEO2469, 2015.
Arévalo-Martínez, D. L., Kock, A., Löscher, C. R., Schmitz, R. A., Stramma, L., and Bange, H. W.: Influence of mesoscale eddies on the distribution of nitrous oxide in the eastern tropical South Pacific, Biogeosciences, 13, 1105–1118, https://doi.org/10.5194/bg-13-1105-2016, 2016.
Babbin, A. R., Keil, R. G., Devol, A. H., and Ward, B. B.: Organic matter
stoichiometry, flux, and oxygen control nitrogen loss in the ocean,
Science, 344, 406–408, https://doi.org/10.1126/science.1248364, 2014.
Babbin, A. R., Bianchi, D., Jayakumar, A., and Ward, B. B.: Rapid nitrous
oxide cycling in the suboxic ocean, Science, 348, 1127–1129,
https://doi.org/10.1126/science.aaa8380, 2015.
Bakker, D. C. E., Bange, H. W., Gruber, N., Johannessen, T., Upstill-Goddard, R. C., Borges, A. V., Delille, B., Löscher, C. R., Naqvi, W. A., Omar, A. M., and Santana-Casiano, J. M.: Air-Sea Interactions of Natural Long-Lived Greenhouse Gases (CO2, N2O, CH4) in a Changing Climate, in: Ocean-Atmosphere Interactions of Gases and Particles, 55–112, Springer, Berlin, Heidelberg, 2014.
Bange, H. W.: Gaseous nitrogen compounds (NO, N2O, N2, NH3)
in the ocean, in: Nitrogen in the marine environment, 51–94, Elsevier,
Amsterdam, 2008.
Battaglia, G. and Joos, F.: Marine N2O Emissions From Nitrification and
Denitrification Constrained by Modern Observations and Projected in
Multimillennial Global Warming Simulations, Global Biogeochem. Cy.,
32, 92–121, https://doi.org/10.1002/2017GB005671, 2018.
Beman, J. M., Chow, C., King, A. L., Feng, Y., and Fuhrman, J. A.: Global
declines in oceanic nitrification rates as a consequence of ocean
acidification, P. Natl. Acad. Sci. USA, 108, 208–213,
https://doi.org/10.1073/pnas.1011053108, 2011.
Bertagnolli, A. D. and Ulloa, O.: Hydrography shapes community composition
and diversity of amoA-containing Thaumarchaeota in the coastal waters off
central Chile, Env. Microb. Rep., 9, 717–728,
https://doi.org/10.1111/1758-2229.12579, 2017.
Biller, S. J., Mosier, A. C., Wells, G. F., and Francis, C. A.: Global
Biodiversity of Aquatic Ammonia-Oxidizing Archaea is Partitioned by Habitat,
Front. Microbiol., 3, 252, https://doi.org/10.3389/fmicb.2012.00252, 2012.
Blum, J. M., Su, Q., Ma, Y., Valverde-Pérez, B., Domingo-Félez, C.,
Jensen, M. M., and Smets, B. F.: The pH dependency of N-converting enzymatic
processes, pathways and microbes: effect on net N2O production,
Environ. Microbiol., 20, 1623–1640, https://doi.org/10.1111/1462-2920.14063, 2018.
Bonaglia, S., Klawonn, I., De Brabandere, L., Deutsch, B., Thamdrup, B., and
Brüchert, V.: Denitrification and DNRA at the Baltic Sea oxic–anoxic
interface: Substrate spectrum and kinetics, Limnol. Oceanogr., 61,
1900–1915, https://doi.org/10.1002/lno.10343, 2016.
Borcard, D., Legendre, P., and Drapeau, P.: Partialling out the Spatial
Component of Ecological Variation, Ecology, 73, 1045–1055,
https://doi.org/10.2307/1940179, 1992.
Bourbonnais, A., Altabet, M. A., Charoenpong, C. N., Larkum, J., Hu, H.,
Bange, H. W., and Stramma, L.: N-loss isotope effects in the Peru oxygen
minimum zone studied using a mesoscale eddy as a natural tracer experiment,
Global Biogeochem. Cy., 29, 793–811, https://doi.org/10.1002/2014GB005001, 2015.
Bourbonnais, A., Letscher, R. T., Bange, H. W., Échevin, V., Larkum, J.,
Mohn, J., Yoshida, N., and Altabet, M. A.: N2O production and
consumption from stable isotopic and concentration data in the Peruvian
coastal upwelling system, Global Biogeochem. Cy., 31, 678–698,
https://doi.org/10.1002/2016GB005567, 2017.
Boyd, P. W., Sherry, N. D., Berges, J. A., Bishop, J. K. B., Calvert, S. E.,
Charette, M. A., Giovannoni, S. J., Goldblatt, R., Harrison, P. J., Moran,
S. B., Roy, S., Soon, M., Strom, S., Thibault, D., Vergin, K. L., Whitney,
F. A., and Wong, C. S.: Transformations of biogenic particulates from the
pelagic to the deep ocean realm, Deep-Sea Res. Pt. II,
46, 2761–2792, https://doi.org/10.1016/S0967-0645(99)00083-1, 1999.
Braker, G., Fesefeldt, A., and Witzel, K. P.: Development of PCR primer
systems for amplification of nitrite reductase genes (nirK and nirS) to
detect denitrifying bacteria in environmental samples, Appl. Environ.
Microb., 64, 3769–3775, 1998.
Breider, F., Yoshikawa, C., Makabe, A., Toyoda, S., Wakita, M., Matsui, Y.,
Kawagucci, S., Fujiki, T., Harada, N., and Yoshida, N.: Response of N2O
production rate to ocean acidification in the western North Pacific, Nat.
Clim. Change, 9, 954–958, https://doi.org/10.1038/s41558-019-0605-7, 2019.
Bristow, L. A., Dalsgaard, T., Tiano, L., Mills, D. B., Bertagnolli, A. D.,
Wright, J. J., Hallam, S. J., Ulloa, O., Canfield, D. E., Revsbech, N. P.,
and Thamdrup, B.: Ammonium and nitrite oxidation at nanomolar oxygen
concentrations in oxygen minimum zone waters, P. Natl. Acad. Sci. USA,
113, 201600359, https://doi.org/10.1073/pnas.1600359113, 2016a.
Bristow, L. A., Callbeck, C. M., Larsen, M., Altabet, M. A., Dekaezemacker,
J., Forth, M., Gauns, M., Glud, R. N., Kuypers, M. M. M., Lavik, G.,
Milucka, J., Naqvi, S. W. A., Pratihary, A., Revsbech, N. P., Thamdrup, B.,
Treusch, A. H., and Canfield, D. E.: N2 production rates limited by
nitrite availability in the Bay of Bengal oxygen minimum zone, Nat. Geosci., 10, 24–29,
https://doi.org/10.1038/ngeo2847, 2016b.
Buitenhuis, E. T., Suntharalingam, P., and Le Quéré, C.: Constraints on global oceanic emissions of N2O from observations and models, Biogeosciences, 15, 2161–2175, https://doi.org/10.5194/bg-15-2161-2018, 2018.
Callbeck, C. M., Lavik, G., Ferdelman, T. G., Fuchs, B., Gruber-Vodicka, H.
R., Hach, P. F., Littmann, S., Schoffelen, N. J., Kalvelage, T., Thomsen,
S., Schunck, H., Löscher, C. R., Schmitz, R. A., and Kuypers, M. M. M.:
Oxygen minimum zone cryptic sulfur cycling sustained by offshore transport
of key sulfur oxidizing bacteria, Nat. Commun., 9, 1–11,
https://doi.org/10.1038/s41467-018-04041-x, 2018.
Capone, D. G. and Hutchins, D. A.: Microbial biogeochemistry of coastal
upwelling regimes in a changing ocean, Nat. Geosci., 6, 711–717,
https://doi.org/10.1038/ngeo1916, 2013.
Caranto, J. D., Lancaster, K. M., Ma, C., Jensen, M. M., Smets, B. F.,
Thamdrup, B., Jayakumar, A., Chang, B. X., Widner, B., Bernhardt, P.,
Mulholland, M. R., Ward, B. B., Sinha, E., Michalak, A. M., Balaji, V.,
Lycus, P., Bøthun, K., Bergaust, L., Shapleigh, J., Bakken, L.,
Frostegård, Å., Massicotte, P., Asmala, E., Stedmon, C., Markager,
S., Caranto, J. D., and Lancaster, K. M.: Nitric oxide is an obligate
bacterial nitrification intermediate produced by hydroxylamine
oxidoreductase, P. Natl. Acad. Sci. USA, 114, 8217–8222,
https://doi.org/10.1073/pnas.1704504114, 2017.
Carini, P., Dupont, C. L., and Santoro, A. E.: Patterns of thaumarchaeal gene
expression in culture and diverse marine environments, Environ. Microbiol., 20, 2112–2124,
https://doi.org/10.1111/1462-2920.14107, 2018.
Carrasco, C., Karstensen, J., and Farias, L.: On the Nitrous Oxide
Accumulation in Intermediate Waters of the Eastern South Pacific Ocean,
Front. Mar. Sci., 4, 24, https://doi.org/10.3389/fmars.2017.00024, 2017.
Casciotti, L. and Ward, B. B.: Phylogenetic analysis of nitric oxide
reductase gene homologues from aerobic ammonia-oxidizing bacteria, FEMS Microbiol. Ecol., 52, 197–205, https://doi.org/10.1016/j.femsec.2004.11.002, 2005.
Casciotti, K. L., Forbes, M., Vedamati, J., Peters, B., Martin, T., and
Mordy, C. W.: Nitrous oxide cycling in the Eastern Tropical South Pacific as
inferred from isotopic and isotopomeric data, Deep-Sea Res. Pt. II, 156, 155–167, https://doi.org/10.1016/J.DSR2.2018.07.014, 2018.
Chang, B. X., Rich, J. R., Jayakumar, A., Naik, H., Pratihary, A., Keil, R.
G., Ward, B. B., and Devol, A. H.: The effect of organic carbon on fixed
nitrogen loss in the eastern tropical South Pacific and Arabian Sea oxygen
deficient zones, Limnol. Oceanogr., 59, 1267–1274,
https://doi.org/10.4319/lo.2014.59.4.1267, 2014.
Charpentier, J., Farias, L., Yoshida, N., Boontanon, N., and Raimbault, P.: Nitrous oxide distribution and its origin in the central and eastern South Pacific Subtropical Gyre, Biogeosciences, 4, 729–741, https://doi.org/10.5194/bg-4-729-2007, 2007.
Chavez, F. P. and Messié, M.: A comparison of Eastern Boundary Upwelling
Ecosystems, Prog. Oceanogr., 83, 80–96,
https://doi.org/10.1016/j.pocean.2009.07.032, 2009.
Codispoti, L. A.: Interesting Times for Marine N2O, Science,
332, 1339–1340, 2010.
Cohen, Y. and Gordon, L.: Nitrous oxide in the oxygen minimum of eastern
tropical North Pacific: evidence for its consumption during denitrification
and possble mechanisms for its production, Deep-Sea Res., 25,
509–524, 1978.
Cornejo, M. and Farías, L.: Following the N2O consumption in the oxygen minimum zone of the eastern South Pacific, Biogeosciences, 9, 3205–3212, https://doi.org/10.5194/bg-9-3205-2012, 2012.
Cornejo D'Ottone, M., Bravo, L., Ramos, M., Pizarro, O., Karstensen, J., Gallegos, M., Correa-Ramirez, M., Silva, N., Farias, L., and Karp-Boss, L.: Biogeochemical characteristics of a long-lived anticyclonic eddy in the eastern South Pacific Ocean, Biogeosciences, 13, 2971–2979, https://doi.org/10.5194/bg-13-2971-2016, 2016.
Crutzen, P. J.: The influence of nitrogen oxides on the atmospheric ozone
content, Q. J. Roy. Meteor. Soc., 96, 320–325,
https://doi.org/10.1002/qj.49709640815, 1970.
Dalsgaard, T., Thamdrup, B., Farías, L., Peter Revsbech, N., and
Revsbech, N. P.: Anammox and denitrification in the oxygen minimum zone of
the eastern South Pacific, Limnol. Oceanogr., 57, 1331–1346,
https://doi.org/10.4319/lo.2012.57.5.1331, 2012.
Dalsgaard, T., Stewart, F. J., Thamdrup, B., De Brabandere, L., Revsbech, N.
P., Ulloa, O., Canfield, D. E., and Delong, E. F.: Oxygen at nanomolar levels
reversibly suppresses process rates and gene expression in anammox and
denitrification in the oxygen minimum zone off northern Chile, MBio, 5,
e01966, https://doi.org/10.1128/mBio.01966-14, 2014.
De Brabandere, L., Canfield, D. E., Dalsgaard, T., Friederich, G. E.,
Revsbech, N. P., Ulloa, O., and Thamdrup, B.: Vertical partitioning of
nitrogen-loss processes across the oxic-anoxic interface of an oceanic
oxygen minimum zone, Environ. Microbiol., 16, 3041–3054,
https://doi.org/10.1111/1462-2920.12255, 2014.
Farías, L., Castro-González, M., Cornejo, M., Charpentier, J. J.,
Faúndez, J., Boontanon, N., and Yoshida, N.: Denitrification and nitrous
oxide cycling within the upper oxycline of the eastern tropical South
Pacific oxygen minimum zone, Limnol. Oceanogr., 54, 132–144,
https://doi.org/10.4319/lo.2009.54.1.0132, 2009.
Frame, C. H. and Casciotti, K. L.: Biogeochemical controls and isotopic signatures of nitrous oxide production by a marine ammonia-oxidizing bacterium, Biogeosciences, 7, 2695–2709, https://doi.org/10.5194/bg-7-2695-2010, 2010.
Frame, C. H., Lau, E., Nolan, E. J., Goepfert, T. J., and Lehmann, M. F.:
Acidification Enhances Hybrid N2O Production Associated with Aquatic
Ammonia-Oxidizing Microorganisms, Front. Microbiol., 7, 2104,
https://doi.org/10.3389/fmicb.2016.02104, 2017.
Francis, C. A., Roberts, K. J., Beman, J. M., Santoro, A. E., and Oakley, B.
B.: Ubiquity and diversity of ammonia-oxidizing archaea in water columns and
sediments of the ocean, P. Natl. Acad. Sci. USA, 102, 14683–14688,
https://doi.org/10.1073/pnas.0506625102, 2005.
Frey, C., Bange, H. W., Achterberg, E. P., Jayakumar, A., Löscher, C. R., Arévalo-Martínez, D. L., León-Palmero, E., Sun, M., Sun, X., Xie, R. C., Oleynik, S., and Ward, B. B.: Nitrous oxide production rates in the eastern tropical South Pacific during METEOR cruise M138, PANGAEA, https://doi.org/10.1594/PANGAEA.914948, 2020.
Ganesh, S., Parris, D. J., Delong, E. F., and Stewart, F. J.: Metagenomic
analysis of size-fractionated picoplankton in a marine oxygen minimum zone,
ISME J., 8, 187–211, https://doi.org/10.1038/ismej.2013.144, 2014.
Goreau, T. J., Kaplan, W. A., Wofsy, S. C., McElroy, M. B., Valois, F. W.,
and Watson, S. W.: Production of and N2O by Nitrifying
Bacteria at Reduced Concentrations of Oxygen, Appl. Environ. Microbiol.,
40, 526–532, 1980.
Goréguès, C., Michotey, V., and Bonin, P.: Isolation of
hydrocarbonoclastic denitrifying bacteria from berre microbial mats,
Ophelia, 58, 263–270, https://doi.org/10.1080/00785236.2004.10410234, 2004.
Granger, J. and Ward, B. B.: Accumulation of nitrogen oxides in
copper-limited cultures of denitrifying bacteria, Limnol. Oceanogr., 48,
313–318, https://doi.org/10.4319/lo.2003.48.1.0313, 2003.
Hamdan, L. J., Coffin, R. B., Sikaroodi, M., Greinert, J., Treude, T., and
Gillevet, P. M.: Ocean currents shape the microbiome of Arctic marine
sediments, ISME J., 7, 685–696, https://doi.org/10.1038/ismej.2012.143, 2012.
Hammer, Ø., Harper, D. A. T., and Ryan, P. D.: PAST: Paleontological
statistics software package, Palaeontol. Electron., 4, 1–9,
https://doi.org/10.1016/j.bcp.2008.05.025, 2001.
Haskell, W. Z., Kadko, D., Hammond, D. E., Knapp, A. N., Prokopenko, M. G.,
Berelson, W. M., and Capone, D. G.: Upwelling velocity and eddy diffusivity
from 7Be measurements used to compare vertical nutrient flux to export POC
flux in the Eastern Tropical South Pacific, Mar. Chem., 168, 140–150,
https://doi.org/10.1016/J.MARCHEM.2014.10.004, 2015.
Hink, L., Nicol, G. W., and Prosser, J. I.: Archaea produce lower yields of
N2O than bacteria during aerobic ammonia oxidation in soil, Environ.
Microbiol., 19, 4829–4837, https://doi.org/10.1111/1462-2920.13282, 2017a.
Hink, L., Lycus, P., Gubry-Rangin, C., Frostegård, Å., Nicol, G. W.,
Prosser, J. I., and Bakken, L. R.: Kinetics of NH3-oxidation, NO-turnover,
N2O-production and electron flow during oxygen depletion in model
bacterial and archaeal ammonia oxidisers, Environ. Microbiol., 19,
4882–4896, https://doi.org/10.1111/1462-2920.13914, 2017b.
Holmes, R. M., Aminot, A., Kérouel, R., Hooker, B. A., and Peterson, B.
J.: A simple and precise method for measuring ammonium in marine and
freshwater ecosystems, Can. J. Fish. Aquat. Sci., 56, 1802–1808, 1999.
Hu, H., Bourbonnais, A., Larkum, J., Bange, H. W., and Altabet, M. A.: Nitrogen cycling in shallow low-oxygen coastal waters off Peru from nitrite and nitrate nitrogen and oxygen isotopes, Biogeosciences, 13, 1453–1468, https://doi.org/10.5194/bg-13-1453-2016, 2016.
Hu, Z., Wessels, H. J. C. T., Alen, T., Jetten, M. S. M., and Kartal, B.:
Nitric oxide-dependent anaerobic ammonium oxidation, Nat. Commun., 10,
1–7, https://doi.org/10.1038/s41467-019-09268-w, 2019.
Hydes, D., Aoyama, M., Aminot, A., Bakker, K., Becker, S., Coverly, S.,
Daniel, A., Dickson, A. G., Grosso, O., Kerouel, R., van Ooijen, J., Sato,
K., Tanhua, T., Woodward, E. M. S., and Zhang, J. Z.: Determination of
dissolved nutrients (N, P, Si) in seawater with high precision and
inter-comparability using gas-segmented continuous flow analysers, Go-sh.
Repeat Hydrogr. Man. IOCCP Rep., 134, 1–87, 2010.
IPCC: Climate Change 2013: The Physical Science Basis. Contribution of
Working Group I to the Fifth Assessment Report of the Intergovernmental
Panel on Climate Change, Cambridge University Press, Cambridge UK and New
York, USA, 2013.
Jayakumar, A., Peng, X., and Ward, B.: Community composition of bacteria
involved in fixed nitrogen loss in the water column of two major oxygen
minimum zones in the ocean, Aquat. Microb. Ecol., 70, 245–259,
https://doi.org/10.3354/ame01654, 2013.
Jayakumar, D. A., Naqvi, S. W. A., and Ward, B. B.: Distribution and relative
quantification of key genes involved in fixed nitrogen loss from the Arabian
Sea oxygen minimum zone, Indian Ocean Biogeochemcal Process. Ecol. Var., 185,
187–203, 2009.
Jebaraj, C. S., Forster, D., Kauff, F., and Stoeck, T.: Molecular Diversity
of Fungi from Marine Oxygen-Deficient Environments (ODEs), 189–208,
Springer, Berlin, Heidelberg, 2012.
Ji, Q., Babbin, A. R., Peng, X., Bowen, J. L., Ward, B. B., and Ji, Q.:
Nitrogen substrate–dependent nitrous oxide cycling in salt marsh sediments,
J. Mar. Res., 7373, 71–92, 2015a.
Ji, Q., Babbin, A. R., Jayakumar, A., and Ward, B. B.: Nitrous oxide
production by nitrification and denitrification in the Eastern Tropical
South Pacific oxygen minimum zone, Geophys. Res. Lett., 42, 10755–10764,
https://doi.org/10.1002/2015GL066853, 2015b.
Ji, Q., Buitenhuis, E., Suntharalingam, P., Sarmiento, J. L., and Ward, B.
B.: Global nitrous oxide production determined by oxygen sensitivity of
nitrification and denitrification, Global Biogeochem. Cy., 32, 1790–1802, https://doi.org/10.1029/2018GB005887, 2018a.
Ji, Q., Frey, C., Sun, X., Jackson, M., Lee, Y.-S., Jayakumar, A., Cornwell, J. C., and Ward, B. B.: Nitrogen and oxygen availabilities control water column nitrous oxide production during seasonal anoxia in the Chesapeake Bay, Biogeosciences, 15, 6127–6138, https://doi.org/10.5194/bg-15-6127-2018, 2018b.
Johnston, H.: Reduction of Stratospheric Ozone by Nitrogen.Oxide Catalysts
from Supersonic Transport Exhaust, Science, 173, 517–522, 1971.
Kalvelage, T., Jensen, M. M., Contreras, S., Revsbech, N. P., Lam, P.,
Günter, M., LaRoche, J., Lavik, G., and Kuypers, M. M. M. M.: Oxygen
sensitivity of anammox and coupled N-cycle processes in oxygen minimum
zones, PLoS One, 6, e29299, https://doi.org/10.1371/journal.pone.0029299, 2011.
Kalvelage, T., Lavik, G., Jensen, M. M., Revsbech, N. P., Loescher, C.,
Schunck, H., Desai, D. K., Hauss, H., Kiko, R., Holtappels, M., LaRoche, J.,
Schmitz, R. A., Graco, M. I., and Kuypers, M. M. M.: Aerobic Microbial
Respiration In Oceanic Oxygen Minimum Zones, PLoS One, 10, e0133526,
https://doi.org/10.1371/journal.pone.0133526, 2015.
Kartal, B., Kuypers, M. M. M., Lavik, G., Schalk, J., Op den Camp, H. J. M.,
Jetten, M. S. M., and Strous, M.: Anammox bacteria disguised as denitrifiers:
nitrate reduction to dinitrogen gas via nitrite and ammonium, Environ.
Microbiol., 9, 635–642, https://doi.org/10.1111/j.1462-2920.2006.01183.x, 2007.
Klawonn, I., Bonaglia, S., Whitehouse, M. J., Littmann, S., Tienken, D.,
Kuypers, M. M. M., Brüchert, V., and Ploug, H.: Untangling hidden
nutrient dynamics: rapid ammonium cycling and single-cell ammonium
assimilation in marine plankton communities, ISME J., 13, 1960–1974,
https://doi.org/10.1038/s41396-019-0386-z, 2019.
Kock, A. and Bange, H. W.: Counting the ocean's greenhouse gas emissions, EOS (Washington. DC), 96, 10–13, https://doi.org/10.1029/2015EO023665, 2015 (data available at: https://memento.geomar.de/de/n2o, last access: 16 April 2020).
Kock, A., Arévalo-Martínez, D. L., Löscher, C. R., and Bange, H. W.: Extreme N2O accumulation in the coastal oxygen minimum zone off Peru, Biogeosciences, 13, 827–840, https://doi.org/10.5194/bg-13-827-2016, 2016.
Kondo, Y. and Moffett, J. W.: Iron redox cycling and subsurface offshore
transport in the eastern tropical South Pacific oxygen minimum zone, Mar.
Chem., 168, 95–103, https://doi.org/10.1016/J.MARCHEM.2014.11.007, 2015.
Körner, H. and Zumft, W. G.: Expression of denitrification enzymes in
response to the dissolved oxygen level and respiratory substrate in
continuous culture of Pseudomonas stutzeri, Appl. Environ. Microb.,
55, 1670–1676, 1989.
Korth, F., Kock, A., Arévalo-Martínez, D. L., and Bange, H. W.:
Hydroxylamine as a Potential Indicator of Nitrification in the Open Ocean,
Geophys. Res. Lett., 46, 2158–2166, https://doi.org/10.1029/2018GL080466, 2019.
Kozlowski, J. A., Stieglmeier, M., Schleper, C., Klotz, M. G., and Stein, L.
Y.: Pathways and key intermediates required for obligate aerobic
ammonia-dependent chemolithotrophy in bacteria and Thaumarchaeota, ISME J.,
10, 1–10, https://doi.org/10.1038/ismej.2016.2, 2016.
Lam, P., Lavik, G., Jensen, M. M., van de Vossenberg, J., Schmid, M., Woebken,
D., Gutiérrez, D., Amann, R., Jetten, M. S. M., and Kuypers, M. M. M.:
Revising the nitrogen cycle in the Peruvian oxygen minimum zone, P. Natl.
Acad. Sci. USA, 106, 4752–4757, 2009.
Lancaster, K. M., Caranto, J. D., Majer, S. H., and Smith, M. A.: Alternative
Bioenergy?: Updates to and Challenges in Nitrification Metalloenzymology,
Joule, 2, 421–441, https://doi.org/10.1016/j.joule.2018.01.018, 2018.
Landolfi, A., Somes, C. J., Koeve, W., Zamora, L. M., and Oschlies, A.:
Oceanic nitrogen cycling and N2O flux perturbations in the
Anthropocene, Global Biogeochem. Cy., 31, 1236–1255,
https://doi.org/10.1002/2017GB005633, 2017.
Larsen, M., Lehner, P., Borisov, S. M., Klimant, I., Fischer, J. P.,
Stewart, F. J., Canfield, D. E., and Glud, R. N.: In situ quantification of
ultra-low O2 concentrations in oxygen minimum zones: Application of
novel optodes, Limnol. Oceanogr.-Meth., 14, 784–800,
https://doi.org/10.1002/lom3.10126, 2016.
Legendre, P. and Legendre, L.: Numerical ecology, Elsevier, New York, NY,
USA, 2012.
Liu, Z., Stewart, G., Kirk Cochran, J., Lee, C., Armstrong, R. A.,
Hirschberg, D. J., Gasser, B., and Miquel, J.-C.: Why do POC concentrations
measured using Niskin bottle collections sometimes differ from those using
in-situ pumps?, Deep-Sea Res. Pt. I, 52, 1324–1344,
https://doi.org/10.1016/J.DSR.2005.02.005, 2005.
Löscher, C. R., Kock, A., Könneke, M., LaRoche, J., Bange, H. W., and Schmitz, R. A.: Production of oceanic nitrous oxide by ammonia-oxidizing archaea, Biogeosciences, 9, 2419–2429, https://doi.org/10.5194/bg-9-2419-2012, 2012.
Löscher, C. R., Bange, H. W., Schmitz, R. A., Callbeck, C. M., Engel, A., Hauss, H., Kanzow, T., Kiko, R., Lavik, G., Loginova, A., Melzner, F., Meyer, J., Neulinger, S. C., Pahlow, M., Riebesell, U., Schunck, H., Thomsen, S., and Wagner, H.: Water column biogeochemistry of oxygen minimum zones in the eastern tropical North Atlantic and eastern tropical South Pacific oceans, Biogeosciences, 13, 3585–3606, https://doi.org/10.5194/bg-13-3585-2016, 2016.
Lutterbeck, H. E., Arévalo-Martínez, D. L., Löscher, C. R., and
Bange, H. W.: Nitric oxide (NO) in the oxygen minimum zone off Peru, Deep-Sea Res. Pt. II, 156, 148–154,
https://doi.org/10.1016/j.dsr2.2017.12.023, 2018.
Martin, J. H., Knauer, G. A., Karl, D. M., and Broenkow, W. W.: VERTEX:
carbon cycling in the northeast Pacific, Deep-Sea Res. Pt. A, 34, 267–285, https://doi.org/10.1016/0198-0149(87)90086-0, 1987.
Martinez-Rey, J., Bopp, L., Gehlen, M., Tagliabue, A., and Gruber, N.: Projections of oceanic N2O emissions in the 21st century using the IPSL Earth system model, Biogeosciences, 12, 4133–4148, https://doi.org/10.5194/bg-12-4133-2015, 2015.
McGillicuddy Jr., D. J., Anderson, L. A., Bates, N. R., Bibby, T., Buesseler, K. O.,
Carlson, C. A., Davis, C. S., Ewart, C., Falkowski, P. G., Goldthwait, S.
A., Hansell, D. A., Jenkins, W. J., Johnson, R., Kosnyrev, V. K., Ledwell,
J. R., Li, Q. P., Siegel, D. A., and Steinberg, D. K.: Eddy/Wind
Interactions Stimulate Extraordinary Mid-Ocean Plankton Blooms, Science, 316, 1021–1026, https://doi.org/10.1126/science.1136256, 2007.
McIlvin, M. R. and Altabet, M. A.: Chemical Conversion of Nitrate and
Nitrite to Nitrous Oxide for Nitrogen and Oxygen Isotopic Analysis in
Freshwater and Seawater, Anal. Chem., 77, 5589–5595, 2005.
McKenney, D. J., Drury, C. F., Findlay, W. I., Mutus, B., McDonnell, T., and
Gajda, C.: Kinetics of denitrification by Pseudomonas fluorescens: Oxygen
effects, Soil Biol. Biochem., 26, 901–908, 1994.
Messié, M. and Chavez, F. P.: Seasonal regulation of primary production
in eastern boundary upwelling systems, Prog. Oceanogr., 134, 1–18,
https://doi.org/10.1016/j.pocean.2014.10.011, 2015.
Mincer, T. J., Church, M. J., Taylor, L. T., Preston, C., Karl, D. M., and
DeLong, E. F.: Quantitative distribution of presumptive archaeal and
bacterial nitrifiers in Monterey Bay and the North Pacific Subtropical Gyre,
Environ. Microbiol., 9, 1162–1175, https://doi.org/10.1111/j.1462-2920.2007.01239.x,
2007.
Murdock, S. A. and Juniper, S. K.: Capturing Compositional Variation in
Denitrifying Communities: a Multiple-Primer Approach That Includes
Epsilonproteobacteria, Appl. Environ. Microb., 83, 1–16, 2017.
NCBI: Gene Expression Omnibus, available at: http://www.ncbi.nlm.nih.gov/geo/, last access: 16 April 2020.
Newell, S. E., Babbin, A. R., Jayakumar, A., and Ward, B. B.: Ammonia
oxidation rates and nitrification in the Arabian Sea, Global Biogeochem.
Cy., 25, 1–10, https://doi.org/10.1029/2010gb003940, 2011.
Nicholls, J. C., Davies, C. A., and Trimmer, M.: High-resolution profiles and
nitrogen isotope tracing reveal a dominant source of nitrous oxide and
multiple pathways of nitrogen gas formation in the central Arabian Sea,
Limnol. Oceanogr., 52, 156–168, https://doi.org/10.4319/lo.2007.52.1.0156, 2007.
Oksanen, J., Blanchet, F. G., Friendly, M., Kindt, R., Legendre, P.,
Mcglinn, D., Minchin, P. R., O'Hara, R. B., Simpson, G. L., Solymos, P.,
Henry, M., Stevens, H., Szoecs, E., and Maintainer, H. W.: Package “vegan”
Title Community Ecology Package, Community Ecol. Packag., 2, 1–297,
2019.
Peng, X., Jayakumar, A., and Ward, B. B.: Community composition of
ammonia-oxidizing archaea from surface and anoxic depths of oceanic oxygen
minimum zones, Front. Microbiol., 4, 1–12,
https://doi.org/10.3389/fmicb.2013.00177, 2013.
Pietri, A., Testor, P., Echevin, V., Chaigneau, A., Mortier, L., Eldin, G.,
Grados, C., Pietri, A., Testor, P., Echevin, V., Chaigneau, A., Mortier, L.,
Eldin, G., and Grados, C.: Finescale Vertical Structure of the Upwelling
System off Southern Peru as Observed from Glider Data, J. Phys. Oceanogr.,
43, 631–646, https://doi.org/10.1175/JPO-D-12-035.1, 2013.
Qin, W., Meinhardt, K. A., Moffett, J. W., Devol, A. H., Armbrust, E. V.,
Ingalls, A. E., and Stahl, D. A.: Influence of Oxygen Availability on the
Activities of Ammonia-oxidizing Archaea, Env. Microbiol. Rep., 9, 250–256,
https://doi.org/10.1111/1758-2229.12525, 2017.
Ravishankara, A. R., Daniel, J. S., and Portmann, R. W.: Nitrous oxide
(N2O): the dominant ozone-depleting substance emitted in the 21st
century, Science, 326, 123–125, https://doi.org/10.1126/science.1176985, 2009.
Richards, T. A., Jones, M. D. M., Leonard, G., and Bass, D.: Marine Fungi:
Their Ecology and Molecular Diversity, Annu. Rev. Mar. Sci., 4, 495–522,
https://doi.org/10.1146/annurev-marine-120710-100802, 2012.
Santoro, A. E. and Casciotti, K. L.: Enrichment and characterization of
ammonia-oxidizing archaea from the open ocean: phylogeny, physiology and
stable isotope fractionation, ISME J., 5, 1796–808,
https://doi.org/10.1038/ismej.2011.58, 2011.
Santoro, A. E., Casciotti, K. L., and Francis, C. A.: Activity, abundance and
diversity of nitrifying archaea and bacteria in the central California
Current, Environ. Microbiol., 12, 1989–2006,
https://doi.org/10.1111/j.1462-2920.2010.02205.x, 2010.
Santoro, A. E., Buchwald, C., McIlvin, M. R., and Casciotti, K. L.: Isotopic
Signature of N2O Produced by Marine Ammonia-Oxidizing Archaea, Science, 333, 1282–1285, https://doi.org/10.1126/science.1208239, 2011.
Santoro, A. E., Dupont, C. L., Richter, R. A., Craig, M. T., Carini, P.,
McIlvin, M. R., Yang, Y., Orsi, W. D., Moran, D. M., and Saito, M. A.:
Genomic and proteomic characterization of “Candidatus Nitrosopelagicus
brevis”: An ammonia-oxidizing archaeon from the open ocean, P. Natl.
Acad. Sci. USA, 112, 1173–1178, https://doi.org/10.1073/PNAS.1416223112, 2015.
Schmidtko, S., Stramma, L., and Visbeck, M.: Decline in global oceanic oxygen
content during the past five decades, Nature, 542, 335–339,
https://doi.org/10.1038/nature21399, 2017.
Schunck, H., Lavik, G., Desai, D. K., Großkopf, T., Kalvelage, T.,
Löscher, C. R., Paulmier, A., Contreras, S., Siegel, H., Holtappels, M.,
Rosenstiel, P., Schilhabel, M. B., Graco, M., Schmitz, R. A., Kuypers, M. M.
M., and Laroche, J.: Giant hydrogen sulfide plume in the oxygen minimum zone
off Peru supports chemolithoautotrophy, PLoS One, 8, e68661,
https://doi.org/10.1371/journal.pone.0068661, 2013.
Segata, N., Izard, J., Waldron, L., Gevers, D., Miropolsky, L., Garrett, W.
S., and Huttenhower, C.: Metagenomic biomarker discovery and explanation,
Genome Biol., 12, R60, https://doi.org/10.1186/gb-2011-12-6-r60, 2011.
Shoun, H., Fushinobu, S., Jiang, L., Kim, S. W., and Wakagi, T.: Fungal
denitrification and nitric oxide reductase cytochrome P450nor, P.
T. Roy. Soc. B, 367, 1186–1194,
https://doi.org/10.1098/rstb.2011.0335, 2012.
Sigman, D. M., Casciotti, K. L., Andreani, M., Barford, C., Galanter, M., and
Böhlke, J. K.: A Bacterial Method for the Nitrogen Isotopic Analysis of
Nitrate in Seawater and Freshwater, Anal. Chem., 73, 4145–4153, 2001.
Stein, L. Y.: Insights into the physiology of ammonia-oxidizing
microorganisms, Curr. Opin. Chem. Biol., 49, 9–15,
https://doi.org/10.1016/J.CBPA.2018.09.003, 2019.
Stewart, F. J., Ulloa, O., and Delong, E. F.: Microbial metatranscriptomics
in a permanent marine oxygen minimum zone, Environ. Microbiol., 14,
23–40, https://doi.org/10.1111/j.1462-2920.2010.02400.x, 2011.
Stewart, F. J., Dalsgaard, T., Young, C. R., Thamdrup, B., Revsbech, N. P.,
Ulloa, O., Canfield, D. E., and Delong, E. F.: Experimental incubations
elicit profound changes in community transcription in OMZ bacterioplankton,
PLoS One, 7, e37118, https://doi.org/10.1371/journal.pone.0037118, 2012.
Stieglmeier, M., Mooshammer, M., Kitzler, B., Wanek, W.,
Zechmeister-Boltenstern, S., Richter, A., and Schleper, C.: Aerobic nitrous
oxide production through N-nitrosating hybrid formation in ammonia-oxidizing
archaea, ISME J., 8, 1135–46, https://doi.org/10.1038/ismej.2013.220, 2014.
Stramma, L., Johnson, G. C., Sprintall, J., and Mohrholz, V.: Expanding
Oxygen-Minimum Zones in the Tropical Oceans, Science, 320,
655–659, 2008.
Stramma, L., Bange, H. W., Czeschel, R., Lorenzo, A., and Frank, M.: On the role of mesoscale eddies for the biological productivity and biogeochemistry in the eastern tropical Pacific Ocean off Peru, Biogeosciences, 10, 7293–7306, https://doi.org/10.5194/bg-10-7293-2013, 2013.
Sun, X., Ji, Q., Jayakumar, A., and Ward, B. B.: Dependence of nitrite
oxidation on nitrite and oxygen in low-oxygen seawater, Geophys. Res. Lett.,
44, 7883–7891, https://doi.org/10.1002/2017GL074355, 2017.
Swan, B. K., Martinez-Garcia, M., Preston, C. M., Sczyrba, A., Woyke, T.,
Lamy, D., Reinthaler, T., Poulton, N. J., Masland, E. D. P., Gomez, M. L.,
Sieracki, M. E., DeLong, E. F., Herndl, G. J., and Stepanauskas, R.:
Potential for chemolithoautotrophy among ubiquitous bacteria lineages in the
dark ocean, Science, 333, 1296–1300,
https://doi.org/10.1126/science.1203690, 2011.
Thamdrup, B. and Dalsgaard, T.: Production of N2 through Anaerobic Ammonium
Oxidation Coupled to Nitrate Reduction in Marine Sediments, Appl. Environ.
Microb., 68, 1312–1318, https://doi.org/10.1128/aem.68.3.1312-1318.2002, 2002.
Tiano, L., Garcia-Robledo, E., Dalsgaard, T., Devol, A. H., Ward, B. B.,
Ulloa, O., Canfield, D. E., and Revsbech, N. P.: Oxygen distribution and
aerobic respiration in the north and south eastern tropical Pacific oxygen
minimum zones, Deep-Sea Res. Pt. I, 94, 173–183,
https://doi.org/10.1016/j.dsr.2014.10.001, 2014.
Torres-Beltrán, M., Mueller, A., Scofield, M., Pachiadaki, M. G.,
Taylor, C., Tyshchenko, K., Michiels, C., Lam, P., Ulloa, O., Jürgens,
K., Hyun, J. H., Edgcomb, V. P., Crowe, S. A., and Hallam, S. J.: Sampling
and processing methods impact microbial community structure and potential
activity in a seasonally anoxic fjord: Saanich inlet, British Columbia,
Front. Mar. Sci., 6, 1–16, https://doi.org/10.3389/fmars.2019.00132, 2019.
Trimmer, M., Chronopoulou, P.-M., Maanoja, S. T., Upstill-Goddard, R. C.,
Kitidis, V., and Purdy, K. J.: Nitrous oxide as a function of oxygen and
archaeal gene abundance in the North Pacific, Nat. Commun., 7, 13451,
https://doi.org/10.1038/ncomms13451, 2016.
Vajrala, N., Martens-Habbena, W., Sayavedra-Soto, L. A., Schauer, A.,
Bottomley, P. J., Stahl, D. A., and Arp, D. J.: Hydroxylamine as an
intermediate in ammonia oxidation by globally abundant marine archaea, P.
Natl. Acad. Sci. USA, 110, 1006–1011, https://doi.org/10.1073/pnas.1214272110, 2013.
Van Der Star, W. R. L., Van De Graaf, M. J., Kartal, B., Picioreanu, C.,
Jetten, M. S. M., and Van Loosdrecht, M. C. M.: Response of anaerobic
ammonium-oxidizing bacteria to hydroxylamine, Appl. Environ. Microb.,
74, 4417–4426, https://doi.org/10.1128/AEM.00042-08, 2008.
Wankel, S. D., Ziebis, W., Buchwald, C., Charoenpong, C., De Beer, Di.,
Dentinger, J., Xu, Z., and Zengler, K.: Evidence for fungal and
chemodenitrification based N2O flux from nitrogen impacted coastal
sediments, Nat. Commun., 8, 1–11, https://doi.org/10.1038/ncomms15595, 2017.
Ward, B. B. and Bouskill, N. J.: The utility of functional gene arrays for
assessing
community composition, relative abundance, and distribution of ammonia-oxidizing bacteria and archaea, in: Methods in
Enzymology, Vol. 496, 373–396, Academic Press Inc., 2011.
Ward, B. B., Tuit, C. B., Jayakumar, A., Rich, J. J., Moffett, J., and Naqvi,
S. W. A.: Organic carbon, and not copper, controls denitrification in oxygen
minimum zones of the ocean, Deep-Sea Res. Pt. I, 55,
1672–1683, https://doi.org/10.1016/j.dsr.2008.07.005, 2008.
Weier, K. L., Doran, J. W., Power, J. F., and Walters, D. T.: Denitrification
and the Dinitrogen/Nitrous Oxide Ratio as Affected by Soil Water, Available
Carbon, and Nitrate, Soil Sci. Soc. Am. J., 57, 66–72,
https://doi.org/10.2136/sssaj1993.03615995005700010013x, 1993.
Weigand, M. A., Foriel, J., Barnett, B., Oleynik, S., and Sigman, D. M.:
Updates to instrumentation and protocols for isotopic analysis of nitrate by
the denitrifier method, Rapid Commun. Mass Sp., 30, 1365–1383,
https://doi.org/10.1002/rcm.7570, 2016.
Wright, J. J., Konwar, K. M., and Hallam, S. J.: Microbial ecology of
expanding oxygen minimum zones, Nat. Rev. Microbiol., 10, 381–394,
https://doi.org/10.1038/nrmicro2778, 2012.
Wuchter, C., Abbas, B., Coolen, M. J. L., Herfort, L., van Bleijswijk, J.,
Timmers, P., Strous, M., Teira, E., Herndl, G. J., Middelburg, J. J.,
Schouten, S., and Sinninghe Damsté, J. S.: Archaeal nitrification in the
ocean, P. Natl. Acad. Sci. USA, 103, 12317–12322,
https://doi.org/10.1073/pnas.0600756103, 2006.
Yang, S., Gruber, N., Long, M. C., and Vogt, M.: High ENSO driven variability
of denitrification and suboxia in the Eastern Pacific Ocean, Global
Biogeochem. Cy., 31, 1470–1487, https://doi.org/10.1002/2016GB005596, 2017.
Yoshida, N.: 15N-depleted N2O as a product of nitrification,
Nature, 335, 528–529, https://doi.org/10.1038/335528a0, 1988.
Zhou, Z., Takaya, N., Sakairi, M. A. C., and Shoun, H.: Oxygen requirement
for denitrification by the fungus Fusarium oxysporum, Arch. Microbiol.,
175, 19–25, https://doi.org/10.1007/s002030000231, 2001.
Zhu-Barker, X., Cavazos, A. R., Ostrom, N. E., Horwath, W. R., and Glass, J.
B.: The importance of abiotic reactions for nitrous oxide production,
Biogeochemistry, 126, 251–267, https://doi.org/10.1007/s10533-015-0166-4, 2015.
Zumft, W. G.: Cell biology and molecular basis of denitrification,
Microbiol. Mol. Biol. Rev., 61, 533–616, 1997.
Short summary
The production of N2O via nitrification and denitrification associated with low-O2 waters is a major source of oceanic N2O. We investigated the regulation and dynamics of these processes with respect to O2 and organic matter inputs. The transcription of the key nitrification gene amoA rapidly responded to changes in O2 and strongly correlated with N2O production rates. N2O production by denitrification was clearly stimulated by organic carbon, implying that its supply controls N2O production.
The production of N2O via nitrification and denitrification associated with low-O2 waters is a...
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