Claire Mahaffey

Amazon River enhances diazotrophy and carbon sequestration in the tropical North Atlantic Ocean

The fresh water discharged by large rivers such as the Amazon is transported hundreds to thousands of kilometers away from the coast by surface plumes. The nutrients delivered by these river plumes contribute to enhanced primary production in the ocean, and the sinking flux of this new production results in carbon sequestration. Here, we report that the Amazon River plume supports N2 fixation far from the mouth and provides important pathways for sequestration of atmospheric CO2 in the western tropical North Atlantic (WTNA). We calculate that the sinking of carbon fixed by diazotrophs in the plume sequesters 1.7 Tmol of C annually, in addition to the sequestration of 0.6 Tmol of C yr−1 of the new production supported by NO3 delivered by the river. These processes revise our current understanding that the tropical North Atlantic is a source of 2.5 Tmol of C to the atmosphere [Mikaloff-Fletcher SE, et al. (2007) Inverse estimates of the oceanic sources and sinks of natural CO2 and the implied oceanic carbon transport. Global Biogeochem Cycles 21, doi:10.1029/2006GB002751]. The enhancement of N2 fixation and consequent C sequestration by tropical rivers appears to be a global phenomenon that is likely to be influenced by anthropogenic activity and climate change.

Mesoscale Eddies Drive Increased Silica Export in the Subtropical Pacific Ocean

Mesoscale eddies may play a critical role in ocean biogeochemistry by increasing nutrient supply, primary production, and efficiency of the biological pump, that is, the ratio of carbon export to primary production in otherwise nutrient-deficient waters. We examined a diatom bloom within a cold-core cyclonic eddy off Hawai`i. Eddy primary production, community biomass, and size composition were markedly enhanced but had little effect on the carbon export ratio. Instead, the system functioned as a selective silica pump. Strong trophic coupling and inefficient organic export may be general characteristics of community perturbation responses in the warm waters of the Pacific Ocean.

Predictable and efficient carbon sequestration in the North Pacific Ocean supported by symbiotic nitrogen fixation

The atmospheric and deep sea reservoirs of carbon dioxide are linked via physical, chemical, and biological processes. The last of these include photosynthesis, particle settling, and organic matter remineralization, and are collectively termed the “biological carbon pump.” Herein, we present results from a 13-y (1992–2004) sediment trap experiment conducted in the permanently oligotrophic North Pacific Subtropical Gyre that document a large, rapid, and predictable summertime (July 15–August 15) pulse in particulate matter export to the deep sea (4,000 m). Peak daily fluxes of particulate matter during the summer export pulse (SEP) average 408, 283, 24.1, 1.1, and 67.5 μmol·m−2·d−1 for total carbon, organic carbon, nitrogen, phosphorus (PP), and biogenic silica, respectively. The SEP is approximately threefold greater than mean wintertime particle fluxes and fuels more efficient carbon sequestration because of low remineralization during downward transit that leads to elevated total carbon/PP and organic carbon/PP particle stoichiometry (371:1 and 250:1, respectively). Our long-term observations suggest that seasonal changes in the microbial assemblage, namely, summertime increases in the biomass and productivity of symbiotic nitrogen-fixing cyanobacteria in association with diatoms, are the main cause of the prominent SEP. The recurrent SEP is enigmatic because it is focused in time despite the absence of any obvious predictable stimulus or habitat condition. We hypothesize that changes in day length (photoperiodism) may be an important environmental cue to initiate aggregation and subsequent export of organic matter to the deep sea.

Alkaline phosphatase activity in the subtropical ocean:insights from nutrient, dust and trace metal addition experiments

Phosphorus is an essential nutrient for all life on earth. In the ocean, the most bioavailable form of phosphorus is inorganic phosphate, but in the extensive subtropical gyres, phosphate concentrations can be chronically low and limit primary productivity and nitrogen fixation. In these regions, organisms produce hydrolytic enzymes, such as alkaline phosphatase (AP), that enable them to utilize the more replete dissolved organic phosphorus (DOP) pool to meet their cellular phosphorus demands. In this study, we synthesized data from 14 published studies and present our own findings from two research cruises (D326 and D361) in the eastern subtropical Atlantic to explore the relationship between AP activity (APA) and nutrients, Saharan dust and trace metals. We found that below a threshold phosphate concentration of ~ 30 nM, APA increased with an inverse hyperbolic relationship with phosphate concentration. Meanwhile, DOP concentrations decreased with enhanced APA, indicating utilization of the DOP pool. We found APA rates were significantly higher in the subtropical Atlantic compared to the subtropical Pacific Ocean, even over the same low phosphate concentration range (0 to 50 nM). While the phosphate concentration may have a first order control on the APA rates, we speculate that other factors influence this basin scale contrast. Using bioassay experiments, we show that the addition of Saharan dust and zinc significantly increased the rate of APA. To our knowledge, our results are the first direct field-based evidence that APA is limited by zinc in the subtropical ocean. Further work is required to explore the relationship between trace metals such as iron and zinc, which are co-factors of phosphohydrolytic enzymes, specifically PhoX and PhoA, respectively, and APA in the ocean.

Evidence for production and lateral transport of dissolved organic phosphorus in the eastern subtropical North Atlantic

The concentration of phosphate and dissolved organic phosphorus (DOP) is chronically low and limits phytoplankton growth in the subtropical North Atlantic relative to other ocean basins. Transport of phosphate and DOP from the productive flanks of the gyre to its interior has been hypothesized as an important phosphorus supply pathway. During a cruise in the eastern Atlantic in spring 2011, the rates of phosphate uptake, alkaline phosphatase activity (APA), and DOP production were measured in the northwest African shelf region, subtropics, and tropics. Rates of DOP production were sixfold higher in the shelf region (43 ± 41 nM d−1) relative to the subtropics (6.9 ± 4.4 nM d−1). In contrast, APA was threefold higher in the subtropics (8.0 ± 7.3 nM d−1), indicative of enhanced DOP utilization, relative to the shelf region (2.6 ± 2.1 nM d−1). Hence, observations suggest net production of DOP in the shelf region and either net consumption of DOP or a near balance in DOP production and consumption in the gyre interior. Eddy-permitting model experiments demonstrate that (i) DOP accounts for over half the total phosphorus in surface waters, (ii) DOP is transported westward from the shelf region by a combination of gyre and eddy circulations, and (iii) advected DOP supports up to 70% of the particle export over much of the subtropical gyre. Our combined observational and modeling study supports the view that the horizontal transport of DOP from the shelf region is an important mechanism supplying phosphorus to the surface subtropical North Atlantic.

High levels of heterogeneity in diazotroph diversity and activity within a putative hotspot for marine nitrogen fixation

Australia’s tropical waters represent predicted ‘hotspots’ for nitrogen (N2) fixation based on empirical and modelled data. However, the identity, activity and ecology of diazotrophs within this region are virtually unknown. By coupling DNA and cDNA sequencing of nitrogenase genes (nifH) with size-fractionated N2 fixation rate measurements, we elucidated diazotroph dynamics across the shelf region of the Arafura and Timor Seas (ATS) and oceanic Coral Sea during Austral spring and winter. During spring, Trichodesmium dominated ATS assemblages, comprising 60% of nifH DNA sequences, while Candidatus Atelocyanobacterium thalassa (UCYN-A) comprised 42% in the Coral Sea. In contrast, during winter the relative abundance of heterotrophic unicellular diazotrophs (δ-proteobacteria and γ-24774A11) increased in both regions, concomitant with a marked decline in UCYN-A sequences, whereby this clade effectively disappeared in the Coral Sea. Conservative estimates of N2 fixation rates ranged from <1 to 91 nmol l−1 day−1, and size fractionation indicated that unicellular organisms dominated N2 fixation during both spring and winter, but average unicellular rates were up to 10-fold higher in winter than in spring. Relative abundances of UCYN-A1 and γ-24774A11 nifH transcripts negatively correlated to silicate and phosphate, suggesting an affinity for oligotrophy. Our results indicate that Australia’s tropical waters are indeed hotspots for N2 fixation and that regional physicochemical characteristics drive differential contributions of cyanobacterial and heterotrophic phylotypes to N2 fixation.

A storm in a shelf sea: Variation in phosphorus distribution and organic matter stoichiometry

Organic matter (OM) plays an important role in productive shelf seas and their contribution to global carbon (C) and nutrient cycles. We investigated the impact of storm mixing on OM dynamics in the seasonally stratified Celtic Sea. After the storm, OM production was decoupled from consumption in the euphotic layer. Over the 15 day study, dissolved OM (DOM) became phosphorus (P) rich relative to C, whereas particulate OM (POM) became P-deplete relative to C. Upward diapycnal phosphate fluxes were accompanied by reciprocal downward mixing of dissolved organic P (DOP) and particulate P (PPhos). Transfer of DOP and PPhos below the thermocline accounts for 22% and 26%, respectively, of the upward phosphate flux. Given the changes in stoichiometry of POM and DOM after the storm, the form in which OM is transferred below the thermocline has important implications for the efficiency of elemental transfer, impacting C cycling and storage in the ocean.

Glider observations of enhanced deep water upwelling at a shelf break canyon: A mechanism for cross-slope carbon and nutrient exchange

Using underwater gliders we have identified canyon driven upwelling across the Celtic Sea shelf-break, in the vicinity of Whittard Canyon. The presence of this upwelling appears to be tied to the direction and strength of the local slope current, which is in itself highly variable. During typical summer time equatorward flow, an unbalanced pressure gradient force and the resulting disruption of geostrophic flow can lead to upwelling along the main axis of two small shelf break canyons. As the slope current reverts to poleward flow, the upwelling stops and the remnants of the upwelled features are mixed into the local shelf water or advected away from the region. The upwelled features are identified by the presence of sub-pycnocline high salinity water on the shelf, and are upwelled from a depth of 300 m on the slope, thus providing a mechanism for the transport of nutrients across the shelf break onto the shelf.