Input Of New Nitrogen Research Articles

An efficient diazotroph‐derived nitrogen transfer pathway in coral reef system

AbstractThe underlying mechanism for sustaining the high productivity of coral reefs in nutrient deplete water, known as the Darwin Paradox, has long been debated. The input of new nitrogen, for example, derived from N2 fixation, and efficient nitrogen recycling are hypothesized as two critical factors in resolving the paradox. However, the source of new nitrogen and its connection to nitrogen recycling in the coral reef system remain poorly understood. We measured N2 fixation rates and investigated the diazotroph community in different components of a tropical coral reef system. Our results show that the reef‐building crustose coralline algae demonstrate a significantly higher N2 fixation rate than the overlying water or the coral holobiont, representing a large source of N2 fixation to the system. The associated diazotroph community was dominated by the non‐cyanobacterial Rhizobiales. 15N2‐based pulse‐chase incubation experiments revealed ~ 50% of the diazotroph‐derived nitrogen was released from the crustose coralline algae holobiont to the overlying water and coral holobiont within 48 h, which was five fold higher than the fraction of nitrogen released from nitrate‐based assimilation by crustose coralline algae. Furthermore, the released diazotroph‐derived nitrogen was rapidly transferred to the coral holobiont through grazing and was subsequently released back to the overlying water as dissolved organic nitrogen, revealing an efficient pathway that connects the new nitrogen input with the nitrogen recycling in the coral reef ecosystem. Our results provide new insights into the sources and fates of new nitrogen in the coral reefs, helping to resolve the Darwin Paradox in this unique ecosystem.

Revealing the transfer pathways of cyanobacterial-fixed N into the boreal forest through the feather-moss microbiome.

Biological N2 fixation in feather-mosses is one of the largest inputs of new nitrogen (N) to boreal forest ecosystems; however, revealing the fate of newly fixed N within the bryosphere (i.e. bryophytes and their associated organisms) remains uncertain. Herein, we combined 15N tracers, high resolution secondary ion mass-spectrometry (NanoSIMS) and a molecular survey of bacterial, fungal and diazotrophic communities, to determine the origin and transfer pathways of newly fixed N2 within feather-moss (Pleurozium schreberi) and its associated microbiome. NanoSIMS images reveal that newly fixed N2, derived from cyanobacteria, is incorporated into moss tissues and associated bacteria, fungi and micro-algae. These images demonstrate that previous assumptions that newly fixed N2 is sequestered into moss tissue and only released by decomposition are not correct. We provide the first empirical evidence of new pathways for N2 fixed in feather-mosses to enter the boreal forest ecosystem (i.e. through its microbiome) and discuss the implications for wider ecosystem function.

Cyanobacterial Diazotroph Distributions in the Western South Atlantic

Inputs of new nitrogen by cyanobacterial diazotrophs are critical to ocean ecosystem structure and function. Relative to other ocean regions, there is a lack of data on the distribution of these microbes in the western South Atlantic. Here, the abundance of six diazotroph phylotypes: Trichodesmium, Crocosphaera, UCYN-A, Richelia associated with Rhizosolenia (Het-1) or Hemiaulus (Het-2), and Calothrix associated with Chaetoceros (Het-3) was measured by quantitative PCR (qPCR) of the nifH gene along a transect extending from the shelf-break to the open ocean along the Vitória-Trindade seamount chain (1200 km). Using nifH gene copies as a proxy for phylotype abundance, Crocosphaera signals were the most abundant, with a broad distribution throughout the study region. Trichodesmium signals were the second most abundant, with the greatest numbers confined to the warmer waters closer to the coast, and a significant positive correlation with temperature. The average signals for the host-associated diazotrophs (UCYN-A, Het-1, and Het-2) were consistently lower than for the other phylotypes. These findings expand measurements of cyanobacterial diazotroph distribution in the western South Atlantic, and provide a new resource to enhance modeling studies focused on patterns of nitrogen fixation in the global ocean.

Warming effect on nitrogen fixation in Mediterranean macrophyte sediments

Abstract. The Mediterranean Sea is warming faster than the global ocean, with important consequences for organisms and biogeochemical cycles. Warming is a major stressor for key marine benthic macrophytes. However, the effect of warming on marine N2 fixation remains unknown, despite the fact that the high productivity of macrophytes in oligotrophic waters is partially sustained by the input of new nitrogen (N) into the system by N2 fixation. Here, we assess the impact of warming on the N2 fixation rates of three key marine macrophytes: Posidonia oceanica, Cymodocea nodosa, and Caulerpa prolifera. We experimentally measured N2 fixation rates in vegetated and bare sediments at temperatures encompassing current summer mean (25 and 27 ∘C), projected summer mean (29 and 31 ∘C), and projected summer maximum (33 ∘C) seawater surface temperatures (SSTs) by the end of the century under a scenario of moderate greenhouse gas emissions. We found that N2 fixation rates in vegetated sediments were 2.8-fold higher than in bare sediments at current summer mean SST, with no differences among macrophytes. Currently, the contribution of N2 fixation to macrophyte productivity could account for up to 7 %, 13.8 %, and 1.8 % of N requirements for P. oceanica, C. nodosa, and C. prolifera, respectively. We show the temperature dependence of sediment N2 fixation rates. However, the thermal response differed for vegetated sediments, in which rates showed an optimum at 31 ∘C followed by a sharp decrease at 33 ∘C, and bare sediments, in which rates increased along the range of the experimental temperatures. The activation energy and Q10 were lower in vegetated than bare sediments, indicating the lower thermal sensitivity of vegetated sediments. The projected warming is expected to increase the contribution of N2 fixation to Mediterranean macrophyte productivity. Therefore, the thermal dependence of N2 fixation might have important consequences for primary production in coastal ecosystems in the context of warming.

Sources and transformations of nitrogen in the South China Sea: insights from nitrogen isotopes

To elucidate the sources and transformations of nitrogen in the South China Sea (SCS), the nitrogen isotopic composition of nitrate ( $${\updelta }^{ 1 5} {\text{N}}_{{{\text{NO}}_{ 3} }}$$ ) was measured in seawater samples from the water column of this marginal sea and the adjacent western North Pacific Ocean (WNP). Comparison of the isotopic signatures from these two locations suggests that the main source of nitrogen into the SCS was nitrate that entered from the WNP through the Luzon Strait. Values of $${\updelta }^{ 1 5} {\text{N}}_{{{\text{NO}}_{ 3} }}$$ were generally lower in the SCS than in the WNP, and the $${\updelta }^{ 1 5} {\text{N}}_{{{\text{NO}}_{ 3} }}$$ maximum observed in the SCS intermediate water was lower than the corresponding WNP maximum. This pattern is attributed to mixing within the SCS in combination with the outflow of SCS intermediate water to the WNP. A mass balance model indicates that atmospherically derived N (a combined input of new nitrogen from marine N2 fixation and atmospheric deposition) supplied approximately 6% of the particulate nitrogen exported from the euphotic zone to the deep SCS. This supply of isotopically light nitrogen cannot, however, explain the low and downward-decreasing δ15N that has been previously observed in sinking particles of the deep SCS. We propose that an alternative explanation might be a downward-increasing ratio of isotopically light NH4 +-N to organic N due to the degradation of organic N within the sinking particles (i.e., relative enrichment of the NH4 +) and also particle incorporation of excreted ammonium from zooplankton.

Importance of salt fingering for new nitrogen supply in the oligotrophic ocean.

The input of new nitrogen into the euphotic zone constrains the export of organic carbon to the deep ocean and thereby the biologically mediated long-term CO2 exchange between the ocean and atmosphere. In low-latitude open-ocean regions, turbulence-driven nitrate diffusion from the ocean's interior and biological fixation of atmospheric N2 are the main sources of new nitrogen for phytoplankton productivity. With measurements across the tropical and subtropical Atlantic, Pacific and Indian oceans, we show that nitrate diffusion (171±190 μmol m−2 d−1) dominates over N2 fixation (9.0±9.4 μmol m−2 d−1) at the time of sampling. Nitrate diffusion mediated by salt fingers is responsible for ca. 20% of the new nitrogen supply in several provinces of the Atlantic and Indian Oceans. Our results indicate that salt finger diffusion should be considered in present and future ocean nitrogen budgets, as it could supply globally 0.23–1.00 Tmol N yr−1 to the euphotic zone.

Strong stimulation of N<sub>2</sub> fixation in oligotrophic Mediterranean Sea: results from dust addition in large in situ mesocosms

Abstract. The response of N2 (dinitrogen) fixation to contrasted (wet and dry) Saharan dust deposition was studied in the framework of the DUNE project (a DUst experiment in a low-Nutrient, low-chlorophyll Ecosystem) during which realistic simulations of dust deposition (10 g m−2) into large mesocosms (52 m3) were performed. Three distinct experimental dust additions were conducted in June 2008 (DUNE-1-P: simulation of a wet deposition, DUNE-1-Q: simulation of a dry deposition) and 2010 (DUNE-2-R: simulation of 2 successive wet depositions) in the northwestern oligotrophic Mediterranean Sea. Here we show that wet and dry dust deposition induced a rapid (24 h or 48 h after dust additions), strong (from 2- to 5.3-fold) and long (at least 4–6 days duration) increase in N2 fixation, indicating that both wet and dry Saharan dust deposition was able to relieve efficiently the nutrient limitation(s) of N2 fixation. This means in particular that N2 fixation activity was not inhibited by the significant input of nitrate associated with the simulated wet deposition (~ 9 mmol NO3− m−2). The input of new nitrogen associated with N2 fixation was negligible relative to the atmospheric NO3− input associated with the dust. The contribution of N2 fixation to primary production was negligible (≤ 1%) before and after dust addition in all experiments, indicating that N2 fixation was a poor contributor to the nitrogen demand for primary production. Despite the stimulation of N2 fixation by dust addition, the rates remained low, and did not significantly change the contribution of N2 fixation to new production since only a maximum contribution of 10% was observed. The response of N2 fixation by diazotrophs and CO2 fixation by the whole phytoplankton community suggests that these metabolic processes were limited or co-limited by different nutrients. With this novel approach, which allows us to study processes as a function of time while atmospheric particles are sinking, we show that new atmospheric nutrients associated with Saharan dust pulses do significantly stimulate N2 fixation in the Mediterranean Sea and that N2 fixation is not a key process in the carbon cycle in such oligotrophic environments.

Nitrogen fixation in restored eelgrass meadows

Biological nitrogen (N2) fixation is the primary input of new nitrogen (N) to marine systems, and is important in meeting the N demands of primary producers. In this study, we deter- mined whether restoration of the eelgrass Zostera marina L. in a shallow coastal bay facilitated increasing rates of N2 fixation as the meadows aged. Rates of N2 fixation were measured in a system that had been devoid of eelgrass following local extinction in the 1930s until restoration by seeding began in 2001. Restored meadows of different ages were compared to nearby bare sediment sites during summer peak metabolism over 2 yr. Nutrient addition by N2 fixation was enhanced as the meadows aged. Rates of N2 fixation in the older (7 to 8 yr old) meadows were 2.7 times more than the younger (2 to 3 yr old) meadows (average 390 and 146 µmol N m �2 d �1 , respectively), and 28 times more than bare sediments (average 14 µmol N m �2 d �1 ). Heterotrophic epiphyte bacteria fixed approximately 90% of the total N2 in Z. marina meadows of both age classes. Both sediment and epiphyte N2 fixation were strongly related to Z. marina density and sediment organic content, suggesting that shoot density increases the positive feedback of plant presence on N2 fixation through the release of organic carbon exudates into the rhizosphere and phyllosphere, and the build up of sediment organic matter also increases. The N provided through fixation represented a large fraction (20.5 to 30%) of the total N demand to support eelgrass aboveground growth during this period of peak summertime production.

Nitrogen fixation in the South Atlantic Gyre and the Benguela Upwelling System

[1] Dinitrogen (N2) fixation is recognized as an important input of new nitrogen (N) to the open ocean gyres, contributing to the export of organic matter from surface waters. However, very little N2-fixation research has focused on the South Atlantic Gyre, where dust deposition of iron (Fe), an important micronutrient for diazotrophs, is seasonally low. Recent modeling efforts suggest that N2-fixation may in fact be closely coupled to, and greatest in, areas of denitrification, as opposed to the oceanic gyres. One of these areas, the Benguela Upwelling System, lies to the east of the South Atlantic Gyre. In this study we show that N2-fixation in surface waters across the South Atlantic Gyre was low overall (<1.5 nmol N l−1 d−1) with highest rates seen in or near the Benguela Upwelling System (up to ∼8 nmol N l−1 d−1). Surface water dissolved Fe (dFe) concentrations were very low in the gyre (∼0.3 nM or lower), while soluble reactive phosphorus (SRP) concentrations were relatively high (∼0.15 μM). N2-fixation rates across the entire sampling area were significantly positively correlated to dFe, but also to SRP and NO3−. Thus, high NO3− concentrations did not exclude N2-fixation in the upwelling region, which provides evidence that N2-fixation may be occurring in previously unrecognized waters, specifically near denitrification zones. However the gene encoding for a nitrogenase component (nifH) was not detected from known diazotrophs at some stations in or near the upwelling where N2-fixation was greatest, suggesting the presence of unknown diazotrophs in these waters.

Constitutive Extracellular Polysaccharide (EPS) Production by Specific Isolates of Crocosphaera watsonii

Unicellular dinitrogen (N2) fixing cyanobacteria have only recently been identified in the ocean and recognized as important contributors to global N2 fixation. The only cultivated representatives of the open ocean unicellular diazotrophs are multiple isolates of Crocosphaera watsonii. Although constituents of the genus are nearly genetically identical, isolates have been described in two size classes, large ∼5 μm and small ∼3 μm cell diameters. We show here that the large size class constitutively produces substantial amounts of extracellular polysaccharides (EPS) during exponential growth, up to 10 times more than is seen in the small size class, and does so under both N2 fixing and non-N2 fixing conditions. The EPS production exceeds the amount produced by larger phytoplankton such as diatoms and coccolithophores by one to two orders of magnitude, is ∼22% of the total particulate organic C in the culture, and is depleted in N compared to cellular material. The large difference in observed EPS production may be accounted for by consistently higher photochemical efficiency of photosystem II in the large (0.5) vs. small (∼0.35) strains. While it is known that Crocosphaera plays an important role in driving the biological carbon (C) pump through the input of new nitrogen (N) to the open ocean, we hypothesize that this species may also contribute directly to the C cycle through the constitutive production of EPS. Indeed, at two stations in the North Pacific Subtropical Gyre, ∼70% of large Crocosphaera cells observed were embedded in EPS. The evolutionary advantage of releasing such large amounts of fixed C is still unknown, but in regions where Crocosphaera can be abundant (i.e., the warm oligotrophic ocean) this material will likely have important biogeochemical consequences.

Interactions between changing pCO2, N2 fixation, and Fe limitation in the marine unicellular cyanobacterium Crocosphaera

We examined the physiological responses of steady‐state iron (Fe)‐replete and Fe‐limited cultures of the biogeochemically critical marine unicellular diazotrophic cyanobacterium Crocosphaera at glacial (19 Pa; 190 ppm), current (39 Pa; 380 ppm), and projected year 2100 (76 Pa; 750 ppm) CO2 levels. Rates of N2 and CO2 fixation and growth increased in step with increasing partial pressure of CO2 (pCO2), but only under Fe‐replete conditions. N2 and carbon fixation rates at 75 Pa CO2 were 1.4‐1.8‐fold and 1.2‐2.0‐fold higher, respectively, relative to those at present day and glacial pCO2 levels. In Fe‐replete cultures, cellular Fe and molybdenum quotas varied threefold and were linearly related to N2 fixation rates and to external pCO2. However, N2 fixation and trace metal quotas were decoupled from pCO2 in Fe‐limited Crocosphaera. Higher CO2 and Fe concentrations both resulted in increased cellular pigment contents and affected photosynthesis vs. irradiance parameters. If these results also apply to natural Crocosphaera populations, anthropogenic CO2 enrichment could substantially increase global oceanic N2 and CO2 fixation, but this effect may be tempered by Fe availability. Possible biogeochemical consequences may include elevated inputs of new nitrogen to the ocean and increased potential for Fe and/or phosphorus limitation in the future high‐CO2 ocean, and feedbacks to atmospheric pCO2 in both the near future and over glacial to interglacial timescales.

DIAZOTROPHIC GROWTH OF THE MARINE CYANOBACTERIUM TRICHODESMIUM IMS101 IN CONTINUOUS CULTURE: EFFECTS OF GROWTH RATE ON N2 -FIXATION RATE, BIOMASS, AND C:N:P STOICHIOMETRY(1).

Trichodesmium N2 fixation has been studied for decades in situ and, recently, in controlled laboratory conditions; yet N2 -fixation rate estimates still vary widely. This variance has made it difficult to accurately estimate the input of new nitrogen (N) by Trichodesmium to the oligotrophic gyres of the world ocean. Field and culture studies demonstrate that trace metal limitation, phosphate availability, the preferential uptake of combined N, light intensity, and temperature may all affect N2 fixation, but the interactions between growth rate and N2 fixation have not been well characterized in this marine diazotroph. To determine the effects of growth rate on N2 fixation, we established phosphorus (P)-limited continuous cultures of Trichodesmium, which we maintained at nine steady-state growth rates ranging from 0.27 to 0.67 d(-1) . As growth rate increased, biomass (measured as particulate N) decreased, and N2 -fixation rate increased linearly. The carbon to nitrogen ratio (C:N) varied from 5.5 to 6.2, with a mean of 5.8 ± 0.2 (mean ± SD, N = 9), and decreased significantly with growth rate. The N:P ratio varied from 23.4 to 45.9, with a mean of 30.5 ± 6.6 (mean ± SD, N = 9), and remained relatively constant over the range of growth rates studied. Relative constancy of C:N:P ratios suggests a tight coupling between the uptake of these three macronutrients and steady-state growth across the range of growth rates. Our work demonstrates that growth rate must be considered when planning studies of the effects of environmental factors on N2 fixation and when modeling the impact of Trichodesmium as a source of new N to oligotrophic regions of the ocean.

Effects of ultraviolet radiation on productivity and nitrogen fixation in the Cyanobacterium, Anabaena sp. (Newton’s strain)

The biological effects of ultraviolet radiation (UVR; 290–400 nm), especially the UV-B (320–400 nm) component of the spectrum, include both direct and indirect effects on many cellular processes. In cyanobacteria both photosynthesis and nitrogen fixation can be affected directly by UVR, and indirectly by UVR through the production of reactive oxygen species (ROS). For the heterocystous cyanobacterium, Anabaena sp. (Newton’s strain), exposure to UVR causes a significant decline in the quantum yields of photosystem II (PSII) fluorescence and maximum productivity despite an increase in UVR absorbing compounds, mycosporine-like amino acids (MAAs), in those cells exposed to UVR. Concurrent with these observations are significant increases in the activities of superoxide dismutase indicative of an increase in the level of oxidative stress in cells exposed to UVR. Additionally, measurements of nitrogenase activity (acetylene reduction) show a significant decrease in cyanobacteria exposed to UVR, which manifests itself as a decrease in cellular nitrogen and an increase in C:N ratios. These results show that these nitrogen-fixing cyanobacteria are particularly sensitive to UVR, both its direct and indirect effects. The effects of UVR reported here add to the increasing evidence that UVR effects on this important group of prokaryotes could affect the input of new nitrogen, and the biogeochemical cycling of this essential macronutrient in terrestrial, marine, and freshwater habitats.

Isotopic heterogeneity and cycling of organic nitrogen in the oligotrophic ocean

We measured the nitrogen (N) isotopic composition (δ15N) of a large set (n = 38) of high‐molecular‐weight (HMW) dissolved organic nitrogen (DON) samples isolated from the tropical and subtropical North Atlantic and Pacific Oceans. The δ15N signature of surface HMW DON is relatively invariable in both oligotrophic basins (4.1 ± 0.6‰ in the Atlantic; 5.4 ± 0.8‰ in the Pacific) and shows little correlation with sources or concentrations of N supporting new production in the euphotic zone. While large variations in δ15N of bulk HMW DON are not apparent, δ15N of proteins isolated from sites with relatively high rates of N2 fixation (&gt;80 µmol N m−2 d−1) were consistently depleted in 15N relative to bulk HMW DON and to proteins isolated from sites where N2 fixation does not routinely occur. This small component of HMW DON appears to be cycling more rapidly than bulk HMW DON and may be indicative of fresh DON contributed by organisms in the surface ocean. Furthermore, δ15N of DNA extracted from the bacterial size fraction (0.2‐0.5 µm) revealed that free‐living bacteria may be an important sink for isotopically depleted N produced during N2 fixation. We suggest that there exists a tight coupling between the production and uptake of DON contributed by diazotrophs (N2 fixers) in regions where N2 fixation provides a major input of new nitrogen.

Denitrification rates and excess nitrogen gas concentrations in the Arabian Sea oxygen deficient zone

Rates of canonical, i.e. heterotrophic, water-column denitrification were measured by 15N incubation techniques at a number of coastal and open ocean stations in the Arabian Sea. Measurements of N 2:Ar gas ratios were also made to obtain independent estimates of N 2 excess resulting from denitrification. Measured denitrification rates ( 15NO 3 − → 15 − 14N 2) at open ocean stations averaged 9.1±1.0 nmol N l −1 d −1 ( n = 15 ), and coastal rates averaged 33.2±12.4 nmol N l −1 d −1 ( n = 18 ). When extrapolated to the entire Arabian Sea, deep measurements within the offshore perennial suboxic zone indicate an overall denitrification rate of 41 Tg N a −1±18 Tg N a −1, which is within the range (10–44 Tg N a −1) of previous estimates for canonical denitrification in the region based on stoichiometric calculations and electron transport system activity. Nitrogen excess gas measurements predict a larger nitrogen anomaly than estimated by classical stoichiometric methods (maximum anomaly=23 μg at N l −1 vs. 13 μg at N l −1, respectively). This mismatch may result from incorrect assumptions of Redfield stoichiometry inherent in the nitrate deficit calculation, inputs of new nitrogen through N-fixation, N 2 contributions from sedimentary denitrification along continental margins, the anammox reaction, and metal catalyzed denitrification reactions. Nevertheless, if denitrification is defined as the conversion of combined nitrogen to a gaseous end product, then the data suggest that denitrification in the Arabian Sea may have been underestimated so far.

Transatlantic Abundance of the N 2 -Fixing Colonial Cyanobacterium Trichodesmium

Colonial diazotrophic cyanobacteria of the genus Trichodesmium are thought to play a significant role in the input of new nitrogen to upper layers of the tropical and subtropical oceanic ecosystems that cover nearly half of Earth's surface. Here we describe results of a transatlantic survey in which a noninvasive underwater digital microscope (the video plankton recorder), was towed across the North Atlantic at 6 meters per second while undulating between the surface and 130 meters. Colony abundance had a basin-scale trend, a clear association with anticyclonic eddies, and was not affected by hurricane-forced mixing. Subsurface abundance was higher than previously reported, which has important implications for the global ocean nitrogen cycle.

Modeling the distribution of Trichodesmium and nitrogen fixation in the Atlantic Ocean

In this paper we use a coupled, 3‐dimensional, biological‐physical model, which includes an explicit, dynamic representation of Trichodesmium, to predict the distribution of Trichodesmium and rates of N2‐fixation in the tropical and subtropical Atlantic Ocean. It is shown that the model reproduces the approximate observed meridional distribution of Trichodesmium in the Atlantic and elevated concentrations in specific coastal and open ocean regions where this organism is known to occur. The model also appears to reproduce the observed seasonality of Trichodesmium populations at higher latitudes (highest concentrations in summer and fall), but this seasonal cycle may be too pronounced at low latitudes. High and persistent Trichodesmium concentrations and rates of N2‐fixation are generated by the model in the Gulf of Guinea off of Africa. This unexpected finding appears to be confirmed by historical measurements. In general, increased Trichodesmium concentrations develop in regions where the mixed layer is relatively thin (resulting in high mean light levels) and dissolved inorganic nitrogen (DIN) concentrations and phytoplankton biomass are low for extended periods of time. The model‐predicted Trichodesmium distributions are therefore very sensitive to the fidelity of the physical model's representation of mixed layer depth variability, and upwelling intensity, and the biological model's estimated DIN and phytoplankton concentrations. The model generates a three‐step successional sequence where (1) high DIN concentrations due to upwelling and/or mixing stimulate phytoplankton growth, followed by (2) Trichodesmium growth after DIN depletion and phytoplankton decline, followed by (3) enhanced phytoplankton growth due to new nitrogen inputs from N2‐fixation. This sequence develops in response to seasonal variations in mixing in the southwestern North Atlantic and in response to upwelling along the coast of Africa and the equator. We interpret this sequence as representing a diatom‐ Trichodesmium‐flagellate succession, which is consistent with observed species successions off of northwest Africa and in the Gulf of Mexico. The results presented in this paper lead us to conclude that our model includes the primary factors that dictate when and where Trichodesmium and N2‐fixation occurs in the Atlantic. Moreover, it appears that our model reproduces some of the major effects that diazotrophically‐derived inputs of new nitrogen have on the pelagic ecosystem.

Nitrogen fixation and nitrogen isotope abundances in zooplankton of the oligotrophic North Atlantic

Deep‐water nitrate is a major reservoir of oceanic combined nitrogen and has long been considered to be the major source of new nitrogen supporting primary production in the oligotrophic ocean. 15N:14N ratios in plankton provide an integrative record of the nitrogen cycle processes at work in the ocean, and near‐surface organic matter in oligotrophic waters like the Sargasso Sea is characterized by an unusually low 15N content relative to average deep‐water nitrate. Herein we show that the low dΔ15N of suspended particles and zooplankton from the tropical North Atlantic cannot arise through isotopic fractionation associated with nutrient uptake and food web processes but are instead consistent with a significant input of new nitrogen to the upper water column by N2 fixation. These results provide direct, integrative evidence that N2 fixation makes a major contribution to the nitrogen budget of the oligotrophic North Atlantic Ocean.

A deterministic model for N2 fixation at stn. ALOHA in the subtropical North Pacific Ocean

Marine N2 fixation by diazotrophic microorganisms is a key process in biogeochemical cycling and yields an important input of new nitrogen into the tropical and subtropical surface ocean. However, it is only poorly accounted for in current numerical models. We present a simple biological model that explicitly includes N2 fixation by diazotrophic phytoplankton. The model employs a mechanistic parameterization of N2 fixation based on physiological responses of Trichodesmium to physical conditions of the environment. The model is conceived to allow shifts in nitrogen versus phosphorus control of the plankton community by resolving the biogeochemical cycles of both elements. Typical N : P ratios were assigned to the different functional groups to capture variations in the N : P stoichiometries of inorganic and organic matter pools. The biological model was coupled to a 1D physical model of the upper ocean. A simulation was performed at stn. ALOHA (22°45′N, 158°W) in the subtropical North Pacific Ocean, where intense blooms of Trichodesmium occurred during the last decade. The model captures essential features of the biological system including the vertical structure and seasonal course of chlorophyll, the seasonal cycle and interannual differences in diazotrophic biomass, the mean vertical particle flux, and an oscillation in the relative importance of nitrogen versus phosphorus. We regard this model as a step towards a mechanistic tool to assess the magnitude of marine N2 fixation and to explore hypotheses on its effect on carbon sequestration from the atmosphere.

Nitrogen fixation by diazotrophic cyanobacteria in the Baltic Sea and transfer of the newly fixed nitrogen to picoplankton organisms

Nitrogen fixation ( 15N 2-tracer method) by filamentous cyanobacteria and the flux of the newly fixed nitrogen into picoplanktonic organisms were investigated during three cruises in the central Baltic Sea in July and August 1995 and July 1996. The diazotrophic cyanobacteria consisted mainly of Aphanizomenon and Nodularia and the picoplankton fraction mainly of Synechococcus, a coccoid nondiazotrophic cyanobacterium. The average N 2-fixation rate measured during the two July studies (3.9 nmol l −1 h −1 in 1995 and 2.4 nmol l −1 h −1 in 1996), was about 13 times higher than in August 1995 (0.3 nmol N 2 l −1 h −1). During the July studies, 5–10% of the total N 2-fixation was found in the picoplankton fraction, but none was found in August. The study shows that in the early stage of a cyanobacterial bloom new nitrogen is provided for the pelagic foodweb through release of recently fixed nitrogen by the diazotrophs, whereas in the late stage the input of new nitrogen occurs mainly through lysing of decaying filamentous cyanobacteria cells. Considerable temporal and local variations in nitrogen fixation found in this study indicate that broader studies in time and location are needed to assess the contribution of N 2-fixation to the nitrogen budget of the Baltic Sea.