Primary production in over half of the world's oceans is limited by fixed nitrogen availability. The main loss term from the fixed nitrogen inventory is the production of dinitrogen gas (N(2)) by heterotrophic denitrification or the more recently discovered autotrophic process, anaerobic ammonia oxidation (anammox). Oceanic oxygen minimum zones (OMZ) are responsible for about 35% of oceanic N(2) production and up to half of that occurs in the Arabian Sea. Although denitrification was long thought to be the only loss term, it has recently been argued that anammox alone is responsible for fixed nitrogen loss in the OMZs. Here we measure denitrification and anammox rates and quantify the abundance of denitrifying and anammox bacteria in the OMZ regions of the Eastern Tropical South Pacific and the Arabian Sea. We find that denitrification rather than anammox dominates the N(2) loss term in the Arabian Sea, the largest and most intense OMZ in the world ocean. In seven of eight experiments in the Arabian Sea denitrification is responsible for 87-99% of the total N(2) production. The dominance of denitrification is reproducible using two independent isotope incubation methods. In contrast, anammox is dominant in the Eastern Tropical South Pacific OMZ, as detected using one of the isotope incubation methods, as previously reported. The abundance of denitrifying bacteria always exceeded that of anammox bacteria by up to 7- and 19-fold in the Eastern Tropical South Pacific and Arabian Sea, respectively. Geographic and temporal variability in carbon supply may be responsible for the different contributions of denitrification and anammox in these two OMZs. The large contribution of denitrification to N(2) loss in the Arabian Sea indicates the global significance of denitrification to the oceanic nitrogen budget.

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Abstract. The Arabian Sea contains one of the three major open-ocean denitrification zones in the world. In addition, pelagic denitrification also occurs over the inner and midshelf off the west coast of India. The major differences between the two environments are highlighted using the available data. The perennial open-ocean system occupies two orders of magnitude larger volume than the seasonal coastal system, however, the latter offers more extreme conditions (greater nitrate consumption leading to complete anoxia). Unlike the open-ocean system, the coastal system seems to have undergone a change (i.e., it has intensified) over the past few decades presumably due to enhanced nutrient loading from land. The two systems also differ from each other with regard to the modes of nitrous oxide (N 2 O) production: In the open-ocean suboxic zone, an accumulation of secondary nitrite (NO

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Abstract.Extensive observations were made during the late Southwest Monsoon of 2004 over the Indian and Omani shelves, and along a transect that extended from the southern coast of Oman to the central west coast of India, tracking the southern leg of the US JGOFS expedition (1994)(1995) in the west. The data are used, in conjunction with satellite-derived data, to investigate long-term trends in chlorophyll and sea surface temperature, indicators of upwelling intensity, and to understand factors that control primary production (PP) in the Arabian Sea, focussing on the role of iron. Our results do not support an intensification of upwelling in the western Arabian Sea, reported to have been caused by the decline in the winter/spring Eurasian snow cover since 1997. We also noticed, for the first time, an unexpected development of high-nutrient, low-chlorophyll condition off the southern Omani coast. This feature, coupled with other characteristics of the system, such as a narrow shelf and relatively low iron concentrations in surface waters, suggest a close similarity between the Omani upwelling system and the Peruvian and California upwelling systems, where PP is limited by iron. Iron limitation of PP may complicate simple relationship between upwelling and PP assumed by previous workers, and contribute to the anomalous offshore occurrence of the most severe oxygen (O 2 ) depletion in the region. Over the much wider Indian shelf, which experiences large-scale bottom water O 2 -depletion in summer, adequate iron supply from reCorrespondence to: S. W. A. Naqvi (naqvi@nio.org) ducing bottom-waters and sediments seems to support moderately high PP; however, such production is restricted to the thin, oxygenated surface layer, probably because of the unsuitability of the O 2 -depleted environment for the growth of oxygenic photosynthesizers.

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The fate of the enormous amount of reactive nitrogen released to the environment by human activities in India is unknown. Here we show occurrence of seasonal stratification and generally low concentrations of dissolved inorganic combined nitrogen, and high molecular nitrogen (N2) to argon ratio, thus suggesting seasonal loss to N2 in anoxic hypolimnia of several dam-reservoirs. However, 15N-experiments yielded low rates of denitrification, anaerobic ammonium oxidation and dissimilatory nitrate reduction to ammonium—except in the presence of methane (CH4) that caused ~12-fold increase in denitrification. While nitrite-dependent anaerobic methanotrophs belonging to the NC10 phylum were present, previously considered aerobic methanotrophs were far more abundant (up to 13.9%) in anoxic hypolimnion. Methane accumulation in anoxic freshwater systems seems to facilitate rapid loss of reactive nitrogen, with generally low production of nitrous oxide (N2O), through widespread coupling between methanotrophy and denitrification, potentially mitigating eutrophication and emissions of CH4 and N2O to the atmosphere.

The Arabian Sea is a productive basin where seasonal upwelling and convective mixing result in high surface nutrient concentrations and widespread algal blooms. The factors controlling primary productivity in the Arabian Sea are of interest because the region contains an intense oxygen minimum zone (OMZ) that is a major sink for nitrate in the ocean. A survey of iron (Fe) distribution and redox chemistry was carried out in 2007 to assess its role in Arabian Sea biogeochemistry, including an investigation of the biological response to Fe additions. Results show that Fe is strongly enriched in the eastern Arabian Sea, associated with the OMZ. Much of the Fe within the OMZ is present as Fe(II), which enhances the residence time and accumulation of Fe. In contrast, Fe concentrations are lower in the western Arabian Sea, separated from the Fe-rich OMZ by a water mass boundary. Consequently, low surface values are associated with monsoon-driven upwelling off the Omani coast. Incubations revealed that primary production during the southwest monsoon was strongly limited by Fe over much of the study area. Incubation of surface waters with Fe resulted in rapid growth of Phaeocystis and up to sixfold increase in chlorophyll. This is the first demonstration of Fe limitation in the Arabian Sea, and the first high resolution zonal survey of dissolved Fe and Fe(II) across the basin. Our findings suggest that models developed to predict the response of the Arabian Sea biogeochemistry to global warming need to consider effects on Fe inputs.