SUMMARY: New data force us to raise previous estimates of oceanic denitrification. Our revised estimate of ~ 450 Tg N yr -1 (Tg = 10 12 g) produces an oceanic fixed N budget with a large deficit ) that can be explained only by positing an ocean that has deviated far from a steady-state, the need for a major upwards revision of fixed N inputs, particularly nitrogen fixation, or both. Oceanic denitrification can be significantly altered by small re-distributions of carbon and dissolved oxygen. Since fixed N is a limiting nutrient, uncompensated changes in denitrification affect the ocean's ability to sequester atmospheric CO 2 via the "biological pump". We have also had to modify our concepts of the oceanic N 2 O regime to take better account of the extremely high N 2 O saturations that can arise in productive, low oxygen waters. Recent results from the western Indian Shelf during a period when hypoxic, suboxic and anoxic waters were present produced a maximum surface N 2 O saturation of > 8000%, a likely consequence of "stop and go" denitrification. The sensitivity of N 2 O production and consumption to small changes in the oceanic dissolved oxygen distribution and to the "spin-up" phase of denitrification suggests that the oceanic source term for N 2 O could change rapidly.

Eutrophication of surface waters and hypoxia in bottom waters has been increasing in many coastal areas, leading to very large depletions of marine life in the affected regions. These areas of high surface productivity and low bottom-water oxygen concentration are caused by increasing runoff of nutrients from land. Although the local ecological and socio-economic effects have received much attention, the potential contribution of increasing hypoxia to global-change phenomena is unknown. Here we report the intensification of one of the largest low-oxygen zones in the ocean, which develops naturally over the western Indian continental shelf during late summer and autumn. We also report the highest accumulations yet observed of hydrogen sulphide (H2S) and nitrous oxide (N2O) in open coastal waters. Increased N2O production is probably caused by the addition of anthropogenic nitrate and its subsequent denitrification, which is favoured by hypoxic conditions. We suggest that a global expansion of hypoxic zones may lead to an increase in marine production and emission of N2O, which, as a potent greenhouse gas, could contribute significantly to the accumulation of radiatively active trace gases in the atmosphere.

Abstract. Coastal hypoxia (defined here as <1.42 ml L −1 ; 62.5 µM; 2 mg L −1 , approx. 30% oxygen saturation) develops seasonally in many estuaries, fjords, and along open coasts as a result of natural upwelling or from anthropogenic eutrophication induced by riverine nutrient inputs. Permanent hypoxia occurs naturally in some isolated seas and marine basins as well as in open slope oxygen minimum zones. Responses of benthos to hypoxia depend on the duration, predictability, and intensity of oxygen depletion and on whether H 2 S is formed. Under suboxic conditions, large mats of filamentous sulfide oxidizing bacteria cover the seabed and consume sulfide. They are hypothesized to provide a detoxified microhabitat for eukaryotic benthic communities. Calcareous foraminiferans and nematodes are particularly tolerant of low oxygen concentrations and may attain high densities and dominance, often in association with microbial mats. When oxygen is sufficient to support metazoans, small, soft-bodied invertebrates (typically annelids), often with short generation times and elaborate branchial structures, predominate. Large taxa are more sensitive than small taxa to hypoxia. Crustaceans and echinoderms are typically more sensitive to hypoxia, with lower oxygen thresholds, than annelids, sipunculans, molluscs and cnidarians.

The anaerobic oxidation of ammonium (anammox) contributes significantly to the global loss of fixed nitrogen and is carried out by a deep branching monophyletic group of bacteria within the phylum Planctomycetes. Various studies have implicated anammox to be the most important process responsible for the nitrogen loss in the marine oxygen minimum zones (OMZs) with a low diversity of marine anammox bacteria. This comprehensive study investigated the anammox bacteria in the suboxic zone of the Black Sea and in three major OMZs (off Namibia, Peru and in the Arabian Sea). The diversity and population composition of anammox bacteria were investigated by both, the 16S rRNA gene sequences and the 16S-23S rRNA internal transcribed spacer (ITS). Our results showed that the anammox bacterial sequences of the investigated samples were all closely related to the Candidatus Scalindua genus. However, a greater microdiversity of marine anammox bacteria than previously assumed was observed. Both phylogenetic markers supported the classification of all sequences in two distinct anammox bacterial phylotypes: Candidatus Scalindua clades 1 and 2. Scalindua 1 could be further divided into four distinct clusters, all comprised of sequences from either the Namibian or the Peruvian OMZ. Scalindua 2 consisted of sequences from the Arabian Sea and the Peruvian OMZ and included one previously published 16S rRNA gene sequence from Lake Tanganyika and one from South China Sea sediment (97.9-99.4% sequence identity). This cluster showed only

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Of the various macro-engineering schemes proposed to mitigate global warming, ocean iron fertilization (OIF) is one that could be started at short notice on relevant scales. It is based on the reasoning that adding trace amounts of iron to iron-limited phytoplankton of the Southern Ocean will lead to blooms, mass sinking of organic matter and ultimately sequestration of significant amounts of atmospheric carbon dioxide (CO 2 ) in the deep sea and sediments. This iron hypothesis, proposed by John Martin in 1990 (Martin 1990 Paleoceanography 5, 1-13), has been tested by five mesoscale experiments that provided strong support for its first condition: stimulation of a diatom bloom accompanied by significant CO 2 drawdown. Nevertheless, a number of arguments pertaining to the fate of bloom biomass, the ratio of iron added to carbon sequestered and various side effects of fertilization, continue to cast doubt on its efficacy. The idea is also unpopular with the public because it is perceived as meddling with nature. However, this apparent consensus against OIF is premature because none of the published experiments were specifically designed to test its second condition pertaining to the fate of iron-induced organic carbon. Furthermore, the arguments on side effects are based on worst-case scenarios. These doubts, formulated as hypotheses, need to be tested in the next generation of OIF experiments. We argue that such experiments, if carried out at appropriate scales and localities, will not only show whether the technique will work, but will also reveal a wealth of insights on the structure and functioning of pelagic ecosystems in general and the krill-based Southern Ocean ecosystem, in particular. The outcomes of current models on the efficacy and side effects of OIF differ widely, so data from adequately designed experiments are urgently needed for realistic parametrization. OIF is likely to boost zooplankton stocks, including krill, which could have a positive effect on recovery of the great whale populations. Negative effects of possible commercialization of OIF can be controlled by the establishment of an international body headed by scientists to supervise and monitor its implementation.

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 mid-shelf 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 (N2O) production: in the open-ocean suboxic zone, an accumulation of secondary nitrite (NO2−) is invariably accompanied by depletion of N2O whereas in the coastal suboxic zone high NO2− and very high N2O concentrations frequently co-occur, indicating, respectively, net consumption and net production of N2O by denitrifiers. The extents of heavier isotope enrichment in the combined nitrate and nitrite (NO3−+NO2−) pool and in N2O in reducing waters appear to be considerably smaller in the coastal region, reflecting more varied sources/sinks and/or different isotopic fractionation factors.