Michael R. Stukel

Variability in diatom contributions to biomass, organic matter production and export across a frontal gradient in the California Current Ecosystem

© 2015. American Geophysical Union. All Rights Reserved. In the offshore waters of Southern California, submesoscale processes associated with fronts may stimulate phytoplankton blooms and lead to biomass shifts at multiple trophic levels. Here we report the results of a study on the cycling of biogenic silica (bSiO 2 ) with estimates of the contributions of diatoms to primary and new production in water masses adjacent to (i.e., coastal or oceanic) and within an offshore front in the Southern California Current Ecosystem (CCE). The coastal and oceanic water were sampled in cyclonic and anticyclonic eddies, respectively, with the frontal water being an interaction region between the eddy types. Concentrations of bSiO 2 varied by 25-fold across the front, with concentrations in frontal waters 20-25% of those in coastal waters. Rates of biogenic silica production spanned an equally large range, with rates within the frontal region that were half those in the coastal regions. Contributions of diatoms to primary and new production were disproportionately higher than their contribution to autotrophic biomass in all areas, ranging from 5-8%, 19-30%, and 32-43% for both processes in the oceanic, frontal and coastal waters, respectively. Across the frontal area, diatoms could account for <1.0%, 6-8%, and 44-72% of organic matter export in the oceanic, frontal and coastal waters, respectively. The results suggest that the regions of frontal interactions between eddies in the southern CCE can account for variability in diatom biomass, productivity and export over very short spatial scales that is comparable to the variability observed across the Pacific basin.

Mesoscale ocean fronts enhance carbon export due to gravitational sinking and subduction

Significance Transport of organic carbon from the sunlit surface ocean to deeper depths drives net oceanic uptake of CO2 from the atmosphere. However, mechanisms that control this carbon export remain poorly constrained, limiting our ability to model and predict future changes in this globally important process. We show that the flux of sinking particles (typically considered the dominant form of downward transport of organic carbon) is twice as high at a frontal system, relative to surrounding waters or to nonfrontal conditions. Furthermore, downward transport by subduction leads to additional carbon export at the front that is similar in magnitude to the sinking flux. Such enhanced C export at episodic and mesoscale features needs to be incorporated into biogeochemical forecast models. Enhanced vertical carbon transport (gravitational sinking and subduction) at mesoscale ocean fronts may explain the demonstrated imbalance of new production and sinking particle export in coastal upwelling ecosystems. Based on flux assessments from 238U:234Th disequilibrium and sediment traps, we found 2 to 3 times higher rates of gravitational particle export near a deep-water front (305 mg C⋅m−2⋅d−1) compared with adjacent water or to mean (nonfrontal) regional conditions. Elevated particle flux at the front was mechanistically linked to Fe-stressed diatoms and high mesozooplankton fecal pellet production. Using a data assimilative regional ocean model fit to measured conditions, we estimate that an additional ∼225 mg C⋅m−2⋅d−1 was exported as subduction of particle-rich water at the front, highlighting a transport mechanism that is not captured by sediment traps and is poorly quantified by most models and in situ measurements. Mesoscale fronts may be responsible for over a quarter of total organic carbon sequestration in the California Current and other coastal upwelling ecosystems.

Enhanced silica ballasting from iron stress sustains carbon export in a frontal zone within the California Current

© 2015. American Geophysical Union. All Rights Reserved. Nutrient dynamics, phytoplankton rate processes, and export were examined in a frontal region between an anticyclone and a pair of cyclones 120 km off the coast in the southern California Current System (sCCS). Low silicic acid: nitrate ratios (Si:N) and high nitrate to iron ratios (N:Fe) characteristic of Fe-limiting conditions in the sCCS were associated with the northern cyclone and with the transition zone between the cyclones and the anticyclone. Phytoplankton growth in low-Si:N, high-N:Fe waters responded strongly to added Fe, confirming growth limitation by Fe of the diatom-dominated phytoplankton community. Low Si:N waters had low biogenic silica content, intermediate productivity, but high export compared to intermediate Si:N waters indicating increased export efficiency under Fe stress. Biogenic silica and particulate organic carbon (POC) export were both high beneath low Si:N waters with biogenic silica export being especially enhanced. This suggests that relatively high POC export from low Si:N waters was supported by silica ballasting from Fe-limited diatoms. Higher POC export efficiency in low Si:N waters may have been further enhanced by lower rates of organic carbon remineralization due to reduced grazing of more heavily armored diatoms growing under Fe stress. The results imply that Fe stress can enhance carbon export, despite lowering productivity, by driving higher export efficiency.

The imbalance of new and export production in the western Antarctic Peninsula, a potentially “leaky” ecosystem

To quantify the balance between new production and vertical nitrogen export of sinking particles, we measured nitrate uptake, net nitrate drawdown, ΔO2/Ar-based net community production, sediment trap flux, and 234Th export at a coastal site near Palmer Station, Antarctica, during the phytoplankton growing season from October 2012 to March 2013. We also measured nitrate uptake and 234Th export throughout the northern western Antarctic Peninsula (WAP) region on a cruise in January 2013. We used a nonsteady state 234Th equation with temporally varying upwelling rates and an irradiance-based phytoplankton production model to correct our export and new production estimates in the complex coastal site near Palmer Station. Results unequivocally showed that nitrate uptake and net community production were significantly greater than the sinking particle export on region-wide spatial scales and season-long temporal scales. At our coastal site, new production (105 ± 17.4 mg N m−2 d−1, mean ± standard error) was 5.3 times greater than vertical nitrogen export (20.4 ± 2.4 mg N m−2 d−1). On the January cruise in the northern WAP, new production (47.9 ± 14.4 mg N m−2 d−1) was 2.4 times greater than export (19.9 ± 1.4 mg N m−2 d−1). Much of this imbalance can be attributed to diffusive losses of particulate nitrogen from the surface ocean due to diapycnal mixing, indicative of a “leaky” WAP ecosystem. If these diffusive losses are common in other systems where new production exceeds export, it may be necessary to revise current estimates of the oceans biological pump.

Using Lagrangian-based process studies to test satellite algorithms of vertical carbon flux in the eastern North Pacific Ocean

Author(s): Stukel, MR; Kahru, M; Benitez-Nelson, CR; Decima, M; Goericke, R; Landry, MR; Ohman, MD | Abstract:

Ocean biogeochemistry modeled with emergent trait-based genomics

Functional ocean biogeography Marine ecosystems are well represented in metagenomic and transcriptomic data. These data are not routinely used to test ecosystem models that explore ocean biogeography or biogeochemistry. Coles et al. built a model in which genes for a range of functions were assigned to different suites of simulated microbes (see the Perspective by Rynearson). Communities emerged from the model with realistic biogeographical and biogeochemical profiles when compared to microbial data collected from the Amazon River plume. However, functional composition trumped the details of taxonomy, and different, coevolving community compositions emerged that provided similar biogeochemical outcomes. Science, this issue p. 1149; see also p. 1129 Results of genomic-based microbe community modeling resemble real-world observations from the Amazon River plume. Marine ecosystem models have advanced to incorporate metabolic pathways discovered with genomic sequencing, but direct comparisons between models and “omics” data are lacking. We developed a model that directly simulates metagenomes and metatranscriptomes for comparison with observations. Model microbes were randomly assigned genes for specialized functions, and communities of 68 species were simulated in the Atlantic Ocean. Unfit organisms were replaced, and the model self-organized to develop community genomes and transcriptomes. Emergent communities from simulations that were initialized with different cohorts of randomly generated microbes all produced realistic vertical and horizontal ocean nutrient, genome, and transcriptome gradients. Thus, the library of gene functions available to the community, rather than the distribution of functions among specific organisms, drove community assembly and biogeochemical gradients in the model ocean.

Stirring up the biological pump: Vertical mixing and carbon export in the Southern Ocean

The biological carbon pump (BCP) transports organic carbon from the surface to the oceans interior via sinking particles, vertically migrating organisms, and passive transport of organic matter by advection and diffusion. While many studies have quantified sinking particles, the magnitude of passive transport remains poorly constrained. In the Southern Ocean weak thermal stratification, strong vertical gradients in particulate organic matter, and weak vertical nitrate gradients suggest that passive transport from the euphotic zone may be particularly important. We compile data from seasonal time-series at a coastal site near Palmer Station, annual regional cruises in the Western Antarctic Peninsula (WAP), cruises throughout the broader Southern Ocean, and SOCCOM autonomous profiling floats to estimate spatial and temporal patterns in vertical gradients of nitrate, particulate nitrogen (PN), and dissolved organic carbon (DOC). Under a steady-state approximation, the ratio of ∂PN/∂z to ∂NO3-/∂z suggests that passive transport of PN may be responsible for removing 46% (37%-58%) of the nitrate introduced into the surface ocean of the WAP (with DOM contributing an additional 3-6%) and for 23% (19%-28%) of the BCP in the broader Southern Ocean. A simple model parameterized with in situ nitrate, PN, and primary production data suggested that passive transport was responsible for 54% of the magnitude of the BCP in the WAP. Our results highlight the potential importance of passive transport (by advection and diffusion) of organic matter in the Southern Ocean, but should only be considered indicative of high passive transport (rather than conclusive evidence) due to our steady-state assumptions.

Spring–summer net community production, new production, particle export and related water column biogeochemical processes in the marginal sea ice zone of the Western Antarctic Peninsula 2012–2014

New production (New P, the rate of net primary production (NPP) supported by exogenously supplied limiting nutrients) and net community production (NCP, gross primary production not consumed by community respiration) are closely related but mechanistically distinct processes. They set the carbon balance in the upper ocean and define an upper limit for export from the system. The relationships, relative magnitudes and variability of New P (from 15NO3– uptake), O2 : argon-based NCP and sinking particle export (based on the 238U : 234Th disequilibrium) are increasingly well documented but still not clearly understood. This is especially true in remote regions such as polar marginal ice zones. Here we present a 3-year dataset of simultaneous measurements made at approximately 50 stations along the Western Antarctic Peninsula (WAP) continental shelf in midsummer (January) 2012–2014. Net seasonal-scale changes in water column inventories (0–150 m) of nitrate and iodide were also estimated at the same stations. The average daily rates based on inventory changes exceeded the shorter-term rate measurements. A major uncertainty in the relative magnitude of the inventory estimates is specifying the start of the growing season following sea-ice retreat. New P and NCP(O2) did not differ significantly. New P and NCP(O2) were significantly greater than sinking particle export from thorium-234. We suggest this is a persistent and systematic imbalance and that other processes such as vertical mixing and advection of suspended particles are important export pathways. This article is part of the theme issue ‘The marine system of the west Antarctic Peninsula: status and strategy for progress in a region of rapid change’.

The Significance of Giant Phaeodarians (Rhizaria) to Biogenic Silica Export in the California Current Ecosystem

In marine ecosystems, many planktonic organisms precipitate biogenic silica (bSiO2) to build silicified skeletons. Among them, giant siliceous rhizarians (>500 μm), including Radiolaria and Phaeodaria, are important contributors to oceanic carbon pools but little is known about their contribution to the marine silica cycle. We report the first analyses of giant phaeodarians to bSiO2 export in the California Current Ecosystem. We measured the silica content of single rhizarian cells ranging in size from 470 to 3,920 μm and developed allometric equations to predict silica content (0.37–43.42 μg Si/cell) from morphometric measurements. Using sediment traps to measure phaeodarian fluxes from the euphotic zone on four cruises, we calculated bSiO2 export produced by two families, the Aulosphaeridae and Castanellidae. Biogenic silica export ranged from <0.01 to 0.63 mmol Si · m 2 · day . These two families alone contributed on average 10% (range 0–80%) of total bSiO2 export from the euphotic zone. Their proportional contributions increased substantially in more oligotrophic regions with lower bSiO2 fluxes. Using the in situ Underwater Vision Profiler 5, we characterized vertical distributions of the giant phaeodarian family Aulosphaeridae to a depth of 500 m and inferred their contribution to bSiO2 export in deeper waters. We found a significant increase of Aulosphaeridae export (<0.01 to 2.82 mmol Si · m 2 · day ) when extended to mesopelagic depths. Using a global data set of in situ profiles, we estimated the significance of Aulosphaeridae to bSiO2 export and revealed that they can act as major exporters of bSiO2 to the mesopelagic zone in various regions.

A new approach for incorporating 15N isotopic data into linear inverse ecosystem models with Markov Chain Monte Carlo sampling

Oceanographic field programs often use δ15N biogeochemical measurements and in situ rate measurements to investigate nitrogen cycling and planktonic ecosystem structure. However, integrative modeling approaches capable of synthesizing these distinct measurement types are lacking. We develop a novel approach for incorporating δ15N isotopic data into existing Markov Chain Monte Carlo (MCMC) random walk methods for solving linear inverse ecosystem models. We test the ability of this approach to recover food web indices (nitrate uptake, nitrogen fixation, zooplankton trophic level, and secondary production) derived from forward models simulating the planktonic ecosystems of the California Current and Amazon River Plume. We show that the MCMC with δ15N approach typically does a better job of recovering ecosystem structure than the standard MCMC or L2 minimum norm (L2MN) approaches, and also outperforms an L2MN with δ15N approach. Furthermore, we find that the MCMC with δ15N approach is robust to the removal of input equations and hence is well suited to typical pelagic ecosystem studies for which the system is usually vastly under-constrained. Our approach is easily extendable for use with δ13C isotopic measurements or variable carbon:nitrogen stoichiometry.