All about nitrite: exploring nitrite sources and sinks in the eastern tropical North Pacific oxygen minimum zone (original) (raw)

Anderson, J. J., Okubo, A., Robbins, A. S., and Richards, F. A.: A model for nitrate distributions in oceanic oxygen minimum zones, Deep-Sea Res. Pt. I, 29, 1113–1140, https://doi.org/10.1016/0198-0149(82)90031-0, 1982.

ASTM international: Standard Guide for Spiking into Aqueous Samples, West Conshohocken, PA, https://compass.astm.org/document/?contentCode=ASTM7CD5810-96R067Cen-US&proxycl=https3A2F2Fsecure.astm.org&fromLogin=true (last access: 9 March 2018.), 2006.

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., Peters, B. D., Mordy, C. W., Widner, B., Casciotti, K. L., and Ward, B. B.: Multiple metabolisms constrain the anaerobic nitrite budget in the Eastern Tropical South Pacific, Global Biogeochem. Cy., 31, 258–271, https://doi.org/10.1002/2016GB005407, 2017.

Babbin, A. R., Buchwald, C., Morel, F. M. M., Wankel, S. D., and Ward, B. B.: Nitrite oxidation exceeds reduction and fixed nitrogen loss in anoxic Pacific waters, Mar. Chem., 224, 103814, https://doi.org/10.1016/J.MARCHEM.2020.103814, 2020.

Bange, H. W., Rixen, T., Johansen, A. M., Siefert, R. L., Ramesh, R., Ittekkot, V., Hoffmann, M. R., and Andreae, M. O.: A revised nitrogen budget of the Arabian Sea, Global Biogeochem. Cy., 14, 1283–1297, https://doi.org/10.1029/1999GB001228, 2000.

Beman, J. M., Leilei Shih, J., and Popp, B. N.: Nitrite oxidation in the upper water column and oxygen minimum zone of the eastern tropical North Pacific Ocean, ISME J., 7, 2192–2205, https://doi.org/10.1038/ismej.2013.96, 2013.

Berg, J. S., Ahmerkamp, S., Pjevac, P., Hausmann, B., Milucka, J., and Kuypers, M. M. M.: How low can they go? Aerobic respiration by microorganisms under apparent anoxia, FEMS Microbiol. Rev., 46, 1–14, https://doi.org/10.1093/FEMSRE/FUAC006, 2022.

Bianchi, D., Babbin, A. R., and Galbraith, E. D.: Enhancement of anammox by the excretion of diel vertical migrators, P. Natl. Acad. Sci. USA, 111, 15653–15658, https://doi.org/10.1073/PNAS.1410790111, 2014.

Bock, E., Koops, H. P., Möller, U. C., and Rudert, M.: A new facultatively nitrite oxidizing bacterium, Nitrobacter vulgaris sp. nov., Arch. Microbiol., 153, 105–110, https://doi.org/10.1007/BF00247805, 1990.

Braman, R. S. and Hendrix, S. A.: Nanogram Nitrite and Nitrate Determination in Environmental and Biological Materials by Vanadium(III) Reduction with Chemiluminescence Detection, Anal. Chem., 61, 2715–2718, 1989.

Brandhorst, W.: Nitrification and Denitrification in the Eastern Tropical North Pacific, ICES J. Mar. Sci., 25, 3–20, 1959.

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, 10601–10606, https://doi.org/10.1073/PNAS.1600359113, 2016.

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, 2017.

Buchwald, C. and Wankel, S. D.: Enzyme-catalyzed isotope equilibriu.: A hypothesis to explain apparent N cycling phenomena in low oxygen environments, Mar. Chem., 244, 104140, https://doi.org/10.1016/J.MARCHEM.2022.104140, 2022.

Buchwald, C., Santoro, A. E., Stanley, R. H. R., and Casciotti, K. L.: Nitrogen cycling in the secondary nitrite maximum of the eastern tropical North Pacific off Costa Rica, Global Biogeochem. Cy., 29, 2061–2081, https://doi.org/10.1002/2015GB005187, 2015.

Bulow, S. E., Rich, J. J., Naik, H. S., Pratihary, A. K., and Ward, B. B.: Denitrification exceeds anammox as a nitrogen loss pathway in the Arabian Sea oxygen minimum zone, Deep-Sea Res. Pt. I, 57, 384–393, https://doi.org/10.1016/J.DSR.2009.10.014, 2010.

Busecke, J. J. M., Resplandy, L., Ditkovsky, S. J., and John, J. G.: Diverging Fates of the Pacific Ocean Oxygen Minimum Zone and Its Core in a Warming World, AGU Adv., 3, e2021AV000470, https://doi.org/10.1029/2021AV000470, 2022.

Casciotti, K. L.: Inverse kinetic isotope fractionation during bacterial nitrite oxidation, Geochim. Cosmochim. Ac., 73, 2061–2076, https://doi.org/10.1016/J.GCA.2008.12.022, 2009.

Casciotti, K. L., Buchwald, C., and McIlvin, M.: Implications of nitrate and nitrite isotopic measurements for the mechanisms of nitrogen cycling in the Peru oxygen deficient zone, Deep-Sea Res. Pt. I, 80, 78–93, https://doi.org/10.1016/J.DSR.2013.05.017, 2013.

Codispoti, L. A. and Packard, T. T.: Denitrification rates in the eastern tropical South Pacific, J. Mar. Res., 38, 453–477, 1980.

Codispoti, L. A. and Richards, F. A.: An analysis of the horizontal regime of denitrification in the eastern tropical North Pacific, Limnol. Oceanogr., 21, 379–388, https://doi.org/10.4319/LO.1976.21.3.0379, 1976.

Codispoti, L. A., Brandes, J. A., Christensen, J. P., Devol, A. H., Naqvi, S. W. A., Paerl, H. W., and Yoshinari, T.: The oceanic fixed nitrogen and nitrous oxide budget.: Moving targets as we enter the anthropocene?, Sci. Mar., 65, 85–105, https://doi.org/10.3989/SCIMAR.2001.65S285, 2001.

Codispoti, L. A., Yoshinari, T., and Devol, A. H.: Suboxic respiration in the oceanic water column, i.: Respiration in Aquatic Ecosystems, edited b.: del Giorgio, P. A. and Williams, P. J. L., Oxford University Press, Oxford, UK ISBN 0 19 852709 8, 225–247, 2005.

Cram, J. A., Fuchsman, C. A., Duffy, M. E., Pretty, J. L., Lekanoff, R. M., Neibauer, J. A., Leung, S. W., Huebert, K. B., Weber, T. S., Bianchi, D., Evans, N., Devol, A. H., Keil, R. G., and McDonnell, A. M. P.: Slow Particle Remineralization, Rather Than Suppressed Disaggregation, Drives Efficient Flux Transfer Through the Eastern Tropical North Pacific Oxygen Deficient Zone, Global Biogeochem. Cy., 36, e2021GB007080, https://doi.org/10.1029/2021GB007080, 2022.

Dalsgaard, T., Canfield, D. E., Peterson, J., Thamdrup, B., and Acuna-Gonzales, J.:N production by anamox in the anoxic water column of Golfo Dulce, Costa Rica, Nature, 422, 606–608, https://doi.org/10.1038/nature01526, 2003.

Dalsgaard, T., Thamdrup, B., Farías, L., 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.

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.

DeVries, T., Deutsch, C., Rafter, P. A., and Primeau, F.: Marine denitrification rates determined from a global 3-D inverse model, Biogeosciences, 10, 2481–2496, https://doi.org/10.5194/bg-10-2481-2013, 2013.

Freitag, A., Rudert, M., and Bock, E.: Growth of Nitrobacter by dissimilatoric nitrate reduction, FEMS Microbiol. Lett., 48, 105–109, https://doi.org/10.1111/J.1574-6968.1987.TB02524.X, 1987.

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.: Regulation of nitrous oxide production in low-oxygen waters off the coast of Peru, Biogeosciences, 17, 2263–2287, https://doi.org/10.5194/bg-17-2263-2020, 2020.

Frey, C., Sun, X., Szemberski, L., Casciotti, K. L., Garcia-Robledo, E., Jayakumar, A., Kelly, C. L., Lehmann, M. F., and Ward, B. B.: Kinetics of nitrous oxide production from ammonia oxidation in the Eastern Tropical North Pacific, Limnol. Oceanogr., 68, 424–438, https://doi.org/10.1002/lno.12283, 2022.

Fuchsman, C. A., Devol, A. H., Saunders, J. K., McKay, C., and Rocap, G.: Niche partitioning of the N cycling microbial community of an offshore oxygen deficient zone, Front. Microbiol., 8, 2384, https://doi.org/10.3389/fmicb.2017.02384, 2017.

Fuchsman, C. A., Palevsky, H. I., Widner, B., Duffy, M., Carlson, M. C. G., Neibauer, J. A., Mulholland, M. R., Keil, R. G., Devol, A. H., and Rocap, G.: Cyanobacteria and cyanophage contributions to carbon and nitrogen cycling in an oligotrophic oxygen-deficient zone, ISME J., 13, 2714–2726, https://doi.org/10.1038/s41396-019-0452-6, 2019.

Füssel, J., Lam, P., Lavik, G., Jensen, M. M., Holtappels, M., Günter, M., and Kuypers, M. M. M.: Nitrite oxidation in the Namibian oxygen minimum zone, ISME J., 6, 1200–1209, https://doi.org/10.1038/ismej.2011.178, 2011.

Füssel, J., Lücker, S., Yilmaz, P., Nowka, B., van Kessel, M. A. H. J., Bourceau, P., Hach, P. F., Littmann, S., Berg, J., Spieck, E., Daims, H., Kuypers, M. M. M., and Lam, P.: Adaptability as the key to success for the ubiquitous marine nitrite oxidizer Nitrococcus, Sci. Adv., 3, https://doi.org/10.1126/sciadv.1700807, 2017.

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, 2013.

Ganesh, S., Bristow, L. A., Larsen, M., Sarode, N., Thamdrup, B., and Stewart, F. J.: Size-fraction partitioning of community gene transcription and nitrogen metabolism in a marine oxygen minimum zone, ISME J., 9, 2682–2696, https://doi.org/10.1038/ismej.2015.44, 2015.

Garcia-Robledo, E., Borisov, S., Klimant, I., and Revsbech, N. P.: Determination of respiration rates in water with sub-micromolar oxygen concentrations, Front. Mar. Sci., 3, 244, https://doi.org/10.3389/FMARS.2016.00244, 2016.

Garcia-Robledo, E., Padilla, C. C., Aldunate, M., Stewart, F. J., Ulloa, O., Paulmier, A., Gregori, G., and Revsbech, N. P.: Cryptic oxygen cycling in anoxic marine zones, P. Natl. Acad. Sci. USA, 114, 8319–8324, https://doi.org/10.1073/PNAS.1619844114, 2017.

Garcia-Robledo, E., Paulmier, A., Borisov, S. M., and Revsbech, N. P.: Sampling in low oxygen aquatic environment.: The deviation from anoxic conditions, Limnol. Oceanogr.-Meth., 19, 733–740, https://doi.org/10.1002/LOM3.10457, 2021.

Granger, J. and Sigman, D. M.: Removal of nitrite with sulfamic acid for nitrate N and O isotope analysis with the denitrifier method, Rapid Commun. Mass Spectrom, 23, 3753–3762, https://doi.org/10.1002/rcm.4307, 2009.

Granger, J. and Wankel, S. D.: Isotopic overprinting of nitrification on denitrification as a ubiquitous and unifying feature of environmental nitrogen cycling, P. Natl. Acad. Sci. USA, 113, E6391–E6400, https://doi.org/10.1073/pnas.1601383113, 2016.

Hamersley, M. R., Lavik, G., Woebken, D., Rattray, J. E., Lam, P., Hopmans, E. C., Sinninghe Damsté, J. S., Krüger, S., Graco, M., Gutiérrez, D., and Kuypers, M. M. M.: Anaerobic ammonium oxidation in the Peruvian oxygen minimum zone, Limnol. Oceanogr., 52, 923–933, https://doi.org/10.4319/LO.2007.52.3.0923, 2007.

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, 1801–1808, https://doi.org/10.1139/f99-128, 1999.

Horak, R. E. A., Ruef, W., Ward, B. B., and Devol, A. H.: Expansion of denitrification and anoxia in the eastern tropical North Pacific from 1972 to 2012, Geophys. Res. Lett., 43, 5252–5260, https://doi.org/10.1002/2016GL068871, 2016.

Ito, T., Minobe, S., Long, M. C., and Deutsch, C.: Upper ocean O2 trend.: 1958–2015, Geophys. Res. Lett., 44, 4214–4223, https://doi.org/10.1002/2017GL073613, 2017.

Jayakumar, A., Wajih, S., Naqvi, A., and Ward, B. B.: Distribution and Relative Quantification of Key Genes Involved in Fixed Nitrogen Loss From the Arabian Sea Oxygen Minimum Zone, i.: Indian Ocean Biogeochem. Process. Ecol. Var., edited by: Wiggert, J. D., Hood, R. R., Wajih, S., Naqvi, A., Brink, K. H., and Smith, S. L., Geophys. Monogr. Ser., 185, https://doi.org/10.1029/2008GM000730, 2009.

Jensen, M. M., Lam, P., Revsbech, N. P., Nagel, B., Gaye, B., Jetten, M. S. M., and Kuypers, M. M. M.: Intensive nitrogen loss over the Omani Shelf due to anammox coupled with dissimilatory nitrite reduction to ammonium, ISME J., 5, 1660–1670, https://doi.org/10.1038/ismej.2011.44, 2011.

Kalvelage, T., Lavik, G., Lam, P., Contreras, S., Arteaga, L., Löscher, C. R., Oschlies, A., Paulmier, A., Stramma, L., and Kuypers, M. M. M.: Nitrogen cycling driven by organic matter export in the South Pacific oxygen minimum zone, Nat. Geosci., 6, 228–234, https://doi.org/10.1038/NGEO1739, 2013.

Keeling, R. F., Körtzinger, A., and Gruber, N.: Ocean deoxygenation in a warming world, Ann. Rev. Mar. Sci., 2, 199–229, https://doi.org/10.1146/annurev.marine.010908.163855, 2010.

Kemeny, P. C., Weigand, M. A., Zhang, R., Carter, B. R., Karsh, K. L., Fawcett, S. E., and Sigman, D. M.: Enzyme-level interconversion of nitrate and nitrite in the fall mixed layer of the Antarctic Ocean, Global Biogeochem. Cy., 30, 1069–1085, https://doi.org/10.1002/2015GB005350, 2016.

Kirstein, K. and Bock, E.: Close genetic relationship between Nitrobacter hamburgensis nitrite oxidoreductase and Escherichia coli nitrate reductases, Arch. Microbiol., 160, 447–453, https://doi.org/10.1007/BF00245305, 1993.

Koch, H., Lücker, S., Albertsen, M., Kitzinger, K., Herbold, C., Spieck, E., Nielsen, P. H., Wagner, M., and Daims, H.: Expanded metabolic versatility of ubiquitous nitrite-oxidizing bacteria from the genus Nitrospira, P. Natl. Acad. Sci. USA, 112, 11371–11376, https://doi.org/10.1073/PNAS.1506533112, 2015.

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.

Kraft, B., Jehmlich, N., Larsen, M., Bristow, L. A., Könneke, M., Thamdrup, B., and Canfield, D. E.: Oxygen and nitrogen production by an ammonia-oxidizing archaeon, Science, 375, 97–100, https://doi.org/10.1126/science.abe6733, 2022.

Kuenen, J. G.: Anammox bacteria: from discovery to application, Nat. Rev. Microbiol., 6, 320–326, https://doi.org/10.1038/nrmicro1857, 2008.

Kuypers, M. M. M., Lavik, G., Woebken, D., Schmid, M., Fuchs, B. M., Amann, R., Jørgensen, B. B., and Jetten, M. S. M.: Massive nitrogen loss from the Benguela upwelling system through anaerobic ammonium oxidation, P. Natl. Acad. Sci. USA, 102, 6478–6483, https://doi.org/10.1073/PNAS.0502088102, 2005.

Lam, P. and Kuypers, M. M. M.: Microbial Nitrogen Cycling Processes in Oxygen Minimum Zones, Ann. Rev. Mar. Sci., 3, 317–345, https://doi.org/10.1146/annurev-marine-120709-142814, 2011.

Lam, P., Lavik, G., Jensen, M. M., Van Vossenberg, J. De, 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, https://doi.org/10.1073/PNAS.0812444106, 2009.

Lam, P., Jensen, M. M., Kock, A., Lettmann, K. A., Plancherel, Y., Lavik, G., Bange, H. W., and Kuypers, M. M. M.: Origin and fate of the secondary nitrite maximum in the Arabian Sea, Biogeosciences, 8, 1565–1577, https://doi.org/10.5194/bg-8-1565-2011, 2011.

Lehner, P., Larndorfer, C., Garcia-Robledo, E., Larsen, M., Borisov, S. M., Revsbech, N. P., Glud, R. N., Canfield, D. E., and Klimant, I.: LUMOS – A Sensitive and Reliable Optode System for Measuring Dissolved Oxygen in the Nanomolar Range, PLOS ONE, 10, e0128125, https://doi.org/10.1371/JOURNAL.PONE.0128125, 2015.

Lipschultz, F., Wofsy, S. C., Ward, B. B., Codispoti, L. A., Friedrich, G., and Elkins, J. W.: Bacterial transformations of inorganic nitrogen in the oxygen-deficient waters of the Eastern Tropical South Pacific Ocean, Deep-Sea Res. Pt. I, 37, 1513–1541, https://doi.org/10.1016/0198-0149(90)90060-9, 1990.

Lomas, M. W. and Lipschultz, F.: Forming the primary nitrite maximu.: Nitrifiers or phytoplankton?, Limnol. Oceanogr., 51, 2453–2467, https://doi.org/10.4319/LO.2006.51.5.2453, 2006.

Luther, G. W.: The role of one- and two-electron transfer reactions in forming thermodynamically unstable intermediates as barriers in multi-electron redox reactions, Aquat. Geochem., 16, 395–420, https://doi.org/10.1007/S10498-009-9082-3, 2010.

Luther, G. W. and Popp, J. I.: Kinetics of the Abiotic Reduction of Polymeric Manganese Dioxide by Nitrit.: An Anaerobic Nitrification Reaction, Aquat. Geochem., 8, 15–36, https://doi.org/10.1023/A:1020325604920, 2002.

Margolskee, A., Frenzel, H., Emerson, S., and Deutsch, C.: Ventilation Pathways for the North Pacific Oxygen Deficient Zone, Global Biogeochem. Cy., 33, 875–890, https://doi.org/10.1029/2018GB006149, 2019.

Martin, J. H., Knauer, G. A., Karl, D. M., and Broenkow, W. W.: VERTE.: carbon cycling in the northeast Pacific, Deep-Sea Res. Pt. I, 34, 267–285, https://doi.org/10.1016/0198-0149(87)90086-0, 1987.

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, https://doi.org/10.1021/ac050528s, 2005.

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.

Moriyasu, R., Evans, N., Bolster, K. M., Hardisty, D. S., and Moffett, J. W.: The Distribution and Redox Speciation of Iodine in the Eastern Tropical North Pacific Ocean, Global Biogeochem. Cy., 34, e2019GB006302, https://doi.org/10.1029/2019GB006302, 2020.

Naqvi, S. W. A.: Some aspects of the oxygen-deficient conditions and denitrification in the Arabian Sea, J. Mar. Res., 45, 1049–1072, 1987.

Nozaki, Y.: A fresh look at element distribution in the North Pacific Ocean, Eos T. Am. Geophys. Un., 78, 221–221, https://doi.org/10.1029/97EO00148, 1997.

Oshiki, M., Satoh, H., and Okabe, S.: Ecology and physiology of anaerobic ammonium oxidizing bacteria, Environ. Microbiol., 18, 2784–2796, https://doi.org/10.1111/1462-2920.13134, 2016.

Padilla, C. C., Bristow, L. A., Sarode, N., Garcia-Robledo, E., Gómez Ramírez, E., Benson, C. R., Bourbonnais, A., Altabet, M. A., Girguis, P. R., Thamdrup, B., and Stewart, F. J.: NC10 bacteria in marine oxygen minimum zones, ISME J., 10, 2067–2071, https://doi.org/10.1038/ismej.2015.262, 2016.

Peng, X., Fuchsman, C. A., Jayakumar, A., Oleynik, S., Martens-Habbena, W., Devol, A. H., and Ward, B. B.: Ammonia and nitrite oxidation in the Eastern Tropical North Pacific, Global Biogeochem. Cy., 29, 2034–2049, https://doi.org/10.1002/2015GB005278, 2015.

Peng, X., Fuchsman, C. A., Jayakumar, A., Warner, M. J., Devol, A. H., and Ward, B. B.: Revisiting nitrification in the Eastern Tropical South Pacific: A focus on controls, J. Geophys. Res.-Oceans, 121, 1667–1684, https://doi.org/10.1002/2015JC011455, 2016.

Penn, J., Weber, T., and Deutsch, C.: Microbial functional diversity alters the structure and sensitivity of oxygen deficient zones, Geophys. Res. Lett., 43, 9773–9780, https://doi.org/10.1002/2016GL070438, 2016.

Peters, B. D., Babbin, A. R., Lettmann, K. A., Mordy, C. W., Ulloa, O., Ward, B. B., and Casciotti, K. L.: Vertical modeling of the nitrogen cycle in the eastern tropical South Pacific oxygen deficient zone using high-resolution concentration and isotope measurements, Global Biogeochem. Cy., 30, 1661–1681, https://doi.org/10.1002/2016GB005415, 2016.

R: A language and environment for statistical computing,https://www.r-project.org/, last access: 30 June 2022.

Starkenburg, S. R., Larimer, F. W., Stein, L. Y., Klotz, M. G., Chain, P. S. G., Sayavedra-Soto, L. A., Poret-Peterson, A. T., Gentry, M. E., Arp, D. J., Ward, B., and Bottomley, P. J.: Complete genome sequence of Nitrobacter hamburgensis X14 and comparative genomic analysis of species within the genus Nitrobacter, Appl. Environ. Microb., 74, 2852–2863, https://doi.org/10.1128/AEM.02311-07, 2008.

Stramma, L., Johnson, G. C., Sprintall, J., and Mohrholz, V.: Expanding oxygen-minimum zones in the tropical oceans, Science (80-), 320, 655–658, https://doi.org/10.1126/science.1153847, 2008.

Strickland, J. D. H. and Parsons, T. R.: A Practical Handbook of Seawater Analysis, Fisheries Research Board of Canada, Ottawa, 1972.

Strohm, T. O., Griffin, B., Zumft, W. G., and Schink, B.: Growth yields in bacterial denitrification and nitrate ammonification, Appl. Environ. Microb., 73, 1420–1424, https://doi.org/10.1128/AEM.02508-06, 2007.

Strous, M., Heijnen, J. J., Kuenen, J. G., and Jetten, M. S. M.: The sequencing batch reactor as a powerful tool for the study of slowly growing anaerobic ammonium-oxidizing microorganisms, Appl. Microbiol. Biot., 50, 589–596, https://doi.org/10.1007/S002530051340, 1998.

Sun, X. and Ward, B. B.: Novel metagenome-assembled genomes involved in the nitrogen cycle from a Pacific oxygen minimum zone, ISME Commun., 1, 1–5, https://doi.org/10.1038/s43705-021-00030-2, 2021.

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.

Sun, X., Kop, L. F. M., Lau, M. C. Y., Frank, J., Jayakumar, A., Lücker, S., and Ward, B. B.: Uncultured Nitrospina-like species are major nitrite oxidizing bacteria in oxygen minimum zones, ISME J., 13, 2391–2402, https://doi.org/10.1038/s41396-019-0443-7, 2019.

Sun, X., Frey, C., Garcia-Robledo, E., Jayakumar, A., and Ward, B. B.: Microbial niche differentiation explains nitrite oxidation in marine oxygen minimum zones, ISME J., 15, 1317–1329, https://doi.org/10.1038/s41396-020-00852-3, 2021.

Sun, X., Frey, C., and Ward, B. B.: Nitrite Oxidation Across the Full Oxygen Spectrum in the Ocean, Global Biogeochem. Cy., 37, e2022GB007548, https://doi.org/10.1029/2022GB007548, 2023.

Suter, E. A., Scranton, M. I., Chow, S., Stinton, D., Medina Faull, L., and Taylor, G. T.: Niskin bottle sample collection aliases microbial community composition and biogeochemical interpretation, Limnol. Oceanogr., 62, 606–617, https://doi.org/10.1002/LNO.10447, 2017.

Tang, W., Tracey, J. C., Carroll, J., Wallace, E., Lee, J. A., Nathan, L., Sun, X., Jayakumar, A., and Ward, B. B.: Nitrous oxide production in the Chesapeake Bay, Limnol. Oceanogr., 67, 2101–2116, https://doi.org/10.1002/LNO.12191, 2022.

Taylor, B. W., Keep, C. F., Hall, R. O., Koch, B. J., Tronstad, L. M., Flecker, A. S., and Ulseth, A. J.: Improving the fluorometric ammonium metho.: matrix effects, background fluorescence, and standard additions, J. N. Am. Benthol. Soc., 26, 167–177, https://doi.org/10.1899/0887-3593(2007)26[167:ITFAMM]2.0.CO;2, 2007.

Thamdrup, B. and Dalsgaard, T.: The fate of ammonium in anoxic manganese oxide-rich marine sediment, Geochim. Cosmochim. Ac., 64, 4157–4164, https://doi.org/10.1016/S0016-7037(00)00496-8, 2000.

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.

Thamdrup, B., Dalsgaard, T., Jensen, M. M., Ulloa, O., Farías, L., and Escribano, R.: Anaerobic ammonium oxidation in the oxygen-deficient waters off northern Chile, Limnol. Oceanogr., 51, 2145–2156, https://doi.org/10.4319/LO.2006.51.5.2145, 2006.

Tiano, L., Garcia-Robledo, E., Dalsgaard, T., Devol, A. H., Ward, B. B., Ulloa, O., Canfield, D. E., and Peter Revsbech, N.: 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.

Tsementzi, D., Wu, J., Deutsch, S., Nath, S., Rodriguez-R, L. M., Burns, A. S., Ranjan, P., Sarode, N., Malmstrom, R. R., Padilla, C. C., Stone, B. K., Bristow, L. A., Larsen, M., Glass, J. B., Thamdrup, B., Woyke, T., Konstantinidis, K. T., and Stewart, F. J.: SAR11 bacteria linked to ocean anoxia and nitrogen loss, Nature, 536, 179–183, https://doi.org/10.1038/nature19068, 2016.

Ulloa, O., Canfield, D. E., DeLong, E. F., Letelier, R. M., and Stewart, F. J.: Microbial oceanography of anoxic oxygen minimum zones, P. Natl. Acad. Sci. USA, 109, 15996–16003, https://doi.org/10.1073/PNAS.1205009109, 2012.

Van de Leemput, I. A., Veraart, A. J., Dakos, V., De Klein, J. J. M., Strous, M., and Scheffer, M.: Predicting microbial nitrogen pathways from basic principles, Environ. Microbiol., 13, 1477–1487, https://doi.org/10.1111/J.1462-2920.2011.02450.X, 2011.

Vedamati, J., Chan, C., and Moffett, J. W.: Distribution of dissolved manganese in the Peruvian Upwelling and Oxygen Minimum Zone, Geochim. Cosmochim. Ac., 156, 222–240, https://doi.org/10.1016/J.GCA.2014.10.026, 2015.

Wanninkhof, R.: Relationship between wind speed and gas exchange over the ocean, J. Geophys. Res.-Oceans, 97, 7373–7382, https://doi.org/10.1029/92JC00188, 1992.

Ward, B. B., Glover, H. E., and Lipschultz, F.: Chemoautotrophic activity and nitrification in the oxygen minimum zone off Peru, Deep-Sea Res. Pt. I, 36, 1031–1051, https://doi.org/10.1016/0198-0149(89)90076-9, 1989.

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.

Ward, B. B., Devol, A. H., Rich, J. J., Chang, B. X., Bulow, S. E., Naik, H., Pratihary, A., and Jayakumar, A.: Denitrification as the dominant nitrogen loss process in the Arabian Sea, Nature,, 461, 78–81, https://doi.org/10.1038/nature08276, 2009.

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.

Wunderlich, A., Meckenstock, R. U., and Einsiedl, F.: A mixture of nitrite-oxidizing and denitrifying microorganisms affects the _δ_18O of dissolved nitrate during anaerobic microbial denitrification depending on the _δ_18O of ambient water, Geochim. Cosmochim. Ac., 119, 31–45, https://doi.org/10.1016/J.GCA.2013.05.028, 2013.