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WBC Series: Fine-scale biophysical controls on nutrient supply, phytoplankton community structure, and carbon export in western boundary current regions

Posted by mmaheigan

· Friday, November 10th, 2017

Sophie Clayton¹, Peter Gaube¹, Takeyoshi Nagai², Melissa M. Omand³, Makio Honda⁴

1. University of Washington
2. Tokyo University of Marine Science and Technology, Japan
3. University of Rhode Island
4. Japan Agency for Marine-Earth Science and Technology, Japan

Western boundary current (WBC) regions are largely thought to be hotspots of productivity, biodiversity, and carbon export. The distinct biogeographical characteristics of the biomes bordering WBC fronts change abruptly from stable, subtropical waters to highly seasonal subpolar gyres. The large-scale convergence of these distinct water masses brings different ecosystems into close proximity allowing for cross-frontal exchange. Although the strong horizontal density gradient maintains environmental gradients, instabilities lead to the formation of meanders, filaments, and rings that mediate the exchange of physical, chemical, and ecological properties across the front. WBC systems also act as large-scale conduits, transporting tracers over thousands of kilometers. The combination of these local perturbations and the short advective timescale for water parcels passing through the system is likely the driver of the enhanced local productivity, biodiversity, and carbon export observed in these regions. Our understanding of biophysical interactions in the WBCs, however, is limited by the paucity of in situ observations, which concurrently resolve chemical, biological, and physical properties at fine spatial and temporal scales (1-10 km, days). Here, we review the current state of knowledge of fine-scale biophysical interactions in WBC systems, focusing on their impacts on nutrient supply, phytoplankton community structure, and carbon export. We identify knowledge gaps and discuss how advances in observational platforms, sensors, and models will help to improve our understanding of physical-biological-ecological interactions across scales in WBCs.

Mechanisms of nutrient supply

Nutrient supply to the euphotic zone occurs over a range of scales in WBC systems. The Gulf Stream and the Kuroshio have been shown to act as large-scale subsurface nutrient streams, supporting large lateral transports of nutrients within the upper thermocline (Pelegrí and Csanady 1991; Pelegrí et al. 1996; Guo et al. 2012; Guo et al. 2013). The WBCs are effective in transporting nutrients in part because of their strong volume transports, but also because they support anomalously high subsurface nutrient concentrations compared to adjacent waters along the same isopycnals (Pelegrí and Csanady 1991; Nagai and Clayton 2017; Komatsu and Hiroe pers. comm.). It is likely that the Gulf Stream and Kuroshio nutrient streams originate near the southern boundary of the subtropical gyres (Nagai et al. 2015a). Recent studies have suggested that nutrients in the Gulf Stream originate even farther south in the Southern Ocean (Williams et al. 2006; Sarmiento et al. 2004). These subsurface nutrients can then be supplied to the surface through a range of vertical supply mechanisms, fueling productivity in the WBC regions.

We currently lack a mechanistic understanding of how elevated nutrient levels in these “nutrient streams” are maintained, since mesoscale stirring should act to homogenize them. While it is well understood that the deepening of the mixed layer toward subpolar regions (along nutrient stream pathways) can drive a large-scale induction of nutrients to the surface layer (Williams et al., 2006), the detailed mechanisms driving the vertical supply of these nutrients to the surface layer at synoptic time and space scales remain unclear. Recent studies focusing on the oceanic (sub)mesoscale (spatial scales of 1-100 km) are starting to reveal mechanisms driving intermittent vertical exchange of nutrients and organisms in and out of the euphotic zone.

Recent surveys that resolved micro-scale mixing processes in the Kuroshio Extension and the Gulf Stream have reported elevated turbulence in the thermocline, likely a result of near-inertial internal waves (Nagai et al. 2009, 2012, 2015b; Kaneko et al. 2012, Inoue et al. 2010). In the Tokara Strait, upstream of the Kuroshio Extension, where the geostrophic flow passes shallow topography, pronounced turbulent mixing oriented along coherent banded layers below the thermocline was observed and linked to high-vertical wavenumber near-inertial internal waves (Nagai et al. 2017; Tsutsumi et al. 2017). Within the Kuroshio Extension, measurements made by autonomous microstructure floats have revealed vigorous microscale temperature dissipation within and below the Kuroshio thermocline over at least 300 km following the main stream, which was attributed to active double-diffusive convection (Nagai et al. 2015c). Within the surface mixed layer, recent studies have shown that downfront winds over the Kuroshio Extension generate strong turbulent mixing (D’Asaro et al. 2011; Nagai et al. 2012). The influence of fine-scale vertical mixing on nutrient supply was observed during a high-spatial resolution biogeochemical survey across the Kuroshio Extension front, revealing fine-scale “tongues” of elevated nitrate arranged along isopycnals (Figure 1, Clayton et al. 2014). Subsequent modeling work has shown that these nutrient tongues are ubiquitous features along the southern flank of the Kuroshio Extension front, formed by submesoscale surface mixed layer fronts (Nagai and Clayton 2017).

Microscale turbulence, double-diffusive convection, and submesoscale stirring are all processes associated with meso- and submesoscale fronts. The results from the studies mentioned above support the hypothesis that WBCs are an efficient conduit for transporting nutrients, not only over large scales but also more locally on fine scales, as isopycnal transporters, lateral stirrers, and diapycnal suppliers. It is the sum of these transport processes that ultimately fuels the elevated primary production observed in these regions.

Figure 1. Vertical sections of nitrate (μM) observed across the Kuroshio Extension in October 2009. The panels are organized such that they line up with respect to the density structure of the Kuroshio Extension Front. Cyan contour lines show the mixed layer depth (taken from Nagai and Clayton 2017).

Phytoplankton biomass, community structure, and dynamics

WBCs separate regions with markedly different biogeochemical and ecological characteristics. Subpolar gyres are productive, highly seasonal, tend to support ecosystems with higher phytoplankton biomass, and can be dominated by large phytoplankton and zooplankton taxa. Conversely, subtropical gyres are mostly oligotrophic, support lower photoautotrophic biomass, and are not characterized by a strong seasonal cycle. In turn, these subtropical regions tend to support ecosystems that comprise smaller cells and a tightly coupled microbial loop. As boundaries to these diverse regions, WBCs are the main conduit linking the equatorial and polar oceans and their resident plankton communities. Within the frontal zones, mesoscale dynamics act to stir water masses together and can transport ecosystems across the WBC into regions of markedly different physical and biological characteristics. Furthermore, mesoscale eddies can modulate vertical fluxes via the displacement of ispycnals during eddy intensification or eddy-induced Ekman pumping, or generating submesoscale patches of vertical exchange. At these smaller scales, vigorous vertical circulations ¾ with magnitudes reaching 100 m/day ¾ can fertilize the euphotic zone or transport phytoplankton out of the surface layer.

Numerous studies have hypothesized that the combination of large-scale transport, mesoscale stirring and transport, and submesoscale nutrient input leads to both high biodiversity and high population densities. Using remote sensing data, D’Ovidio et al. (2010) showed that mesoscale stirring in the Brazil-Malvinas Confluence Zone brings together communities from very different source regions, driving locally enhanced biodiversity. In a numerical model, in which physical and biological processes can be explicitly separated and quantified, Clayton et al. (2013) showed that high modeled biodiversity in the WBCs was due to a combination of transport and local nutrient enhancements. And finally, in situ taxonomic surveys crossing the Brazil-Malvinas Confluence (Cermeno et al. 2008) and the Kuroshio Extension (Honjo and Okada 1974; Clayton et al, 2017) showed both enhanced biomass and biodiversity associated with the WBC fronts. Beyond these local enhancements, WBCs might play a larger role in setting regional biogeography. Sugie and Suzuki (2017) found a mixture of temperate and subpolar diatom species in the Kuroshio Extension, suggesting that the boundary current might play a key role in setting downstream diatom diversity.

However tantalizing these results are, they remain relatively inconclusive, in part because of their relatively small temporal and spatial scales. Extending existing approaches for assessing phytoplankton community structure, leveraging emerging ‘omics and continuous sampling techniques, larger regions might be surveyed at high taxonomic and spatial resolution. Combining genomic and transcriptomic observations would provide measures of both organism abundance and activity (Hunt et al. 2013), as well as the potential to better define the relative roles of growth and loss processes. With genetically resolved data and appropriate survey strategies, it will be possible to conclusively determine the presence of these biodiversity hotspots. A better characterization and deeper understanding of these regions will provide insight into the long-term and large-scale biodiversity, stability, and function of the global planktonic ecosystem.

Organic carbon export via physical and biological processes

Export, the removal of fixed carbon from the surface ocean, is driven by gravitational particle sinking, active transport, and (sub)mesoscale processes such as eddy-driven subduction. While evidence suggests that WBCs are likely hot spots of biological (Siegel et al. 2014; Honda et al. 2017a) and physical (Omand et al. 2015) export fluxes out of the euphotic zone, only a small handful of studies have explored this. Recent results from sediment trap studies at the Kuroshio Extension Observatory (KEO) mooring, located just south of the Kuroshio Extension, suggest that there is a link between the passage of mesoscale eddies and carbon export (Honda et al. 2017b). They observed that high export events at 5000 m lagged behind the passage of negative (cyclonic) sea surface height anomalies (SSHA) at the mooring by one to two months (Figure 2). In other regions, underway measurements (Stanley et al. 2010) and optical sensors on autonomous platforms (Briggs et al. 2011; Estapa et al. 2013; Estapa et al. 2015; Bishop et al. 2016) have revealed large episodicity in export proxies over timescales of hours to days and spatial scales of 1-10 km.

Figure 2. Time series of ocean temperature in the upper ~550 m (less than 550 dbar) at station KEO between July 2014 and June 2016. The daily data shown in the figure are available on the KEO database. White contour lines show the temporal variability in the daily satellite-based sea surface height anomaly (SSHA). White open bars show the total mass flux (TMF) observed by the time series sediment trap at 5000 m (based on a figure in Honda et al. 2017b).

Another avenue of carbon export from the surface ocean results from grazing and vertical migration. Vertically migrating zooplankton feed near the surface in the dark and evade predation at depth during the day. Fronts generated by WBCs produce gradients in zooplankton communities, both in terms of grazer biomass and species compositions (e.g., Wiebe and Flierl, 1983), and influence the extent and magnitude of diel vertical migrations. Submesoscale variability in zooplankton abundance can be observed readily in echograms collected by active acoustic sensors, but submesoscale variability in zooplankton community structure and dynamics remains difficult to measure. Thus, the nature of this variability remains largely unknown.

Future research directions

Building a better understanding of how physical and biogeochemical dynamics in WBC regions interact relies on observing these systems at the appropriate scales. This is particularly challenging because of the range of scales at play in these systems and the limitation of existing in situ and remote observing platforms and techniques. As has been outlined above, the ecological and biogeochemical environment of WBCs is the result of long range transport from the flanking subtropical and subpolar gyres, as well as local modification by meso- and submesocale physical dynamics in these frontal systems.

Another challenge in disentangling the relationships between physical and biogeochemical processes in WBCs is the difficulty in measuring rates rather than standing stocks. In such dynamic systems, lags in biological responses mean that the changes in standing stocks may not be collocated with the physical process forcing them. Small-scale lateral stirring spatially and temporally decouples net community production and export while secondary circulations contribute to vertical transport. As much as possible, future process studies should include approaches that can explicitly quantify biological rates and physical transport pathways. New platforms are beginning to fill these observational gaps: BGC-Argo floats, autonomous platforms (e.g., Saildrone), high-frequency underway measurements, and continuous cytometers (including imaging cytometers) are all capable of generating high-spatial resolution datasets of biological and chemical properties over large regions. Gliders and profiling platforms (e.g., WireWalker) are making it possible to measure vertical profiles of biogeochemical properties at high frequency. Operating within a Lagrangian framework, while resolving lateral gradients of physical and biogeochemical tracers with ships or autonomous vehicles, may someday allow us to quantitatively partition the observed small-scale variability in biogeochemical tracers between that attributable to biological or physical processes.

References

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The Ross Sea deep microbial community’s role in sequestering CO2

Posted by mmaheigan

· Thursday, November 9th, 2017

Antarctic shelf systems generate the densest waters in the world. These shelf waters are the building blocks of Antarctic Bottom Water, the ocean’s abyssal water mass. These bottom waters have the potential to sequester carbon out of the atmosphere for millennia. One such form of marine carbon is dissolved organic carbon (DOC). DOC is produced in the surface ocean via primary production and is the global ocean’s largest standing stock of reduced carbon.

In a recent study, Bercovici et al (2017) used hydrographic and biogeochemical measurements to assess the mechanism that brings DOC into the shelf waters of the Ross Sea, the shelf system in the Pacific sector of Antarctica. These mechanisms include sinking particles, brine rejection caused by katabatic winds in the Terra Nova Bay polynya, and vertical mixing. This study revealed that DOC is primarily introduced into the deeper shelf waters via convective overturning and deep vertical mixing upon the onset of austral winter. Substantial DOC enrichment of shelf waters suggests that this carbon is exported off the shelf into Antarctic Bottom Water. However, this study finds much of the excess Ross Sea shelf DOC is actually consumed and remineralized to CO₂ by deep microbial communities at the slope of the Ross Sea shelf, ultimately sequestering this carbon into the ocean’s interior.

Physical and biological processes have the potential to introduce carbon into the dense shelf waters (blue) in the Ross Sea. Incoming waters (yellow) are modified from the Southern Ocean’s circumpolar waters. At the onset of winter, cooler temperatures and katabatic winds cause brine rejection. The rejection of brine, sinking particles and vertical mixing are all potential mechanisms for bringing DOC to the dense shelf waters. At the shelf slope, outflowing shelf waters ultimately contribute to Antarctic Bottom Water formation. This research furthers our understanding of global carbon cycling through demonstrating that Antarctic shelf systems have the potential to sequester organic carbon into the abyssal ocean.

Authors:
Sarah K. Bercovici (Rosenstiel School of Marine and Atmospheric Science, University of Miami)
Bruce A. Huber (Lamont Doherty Earth Observatory, Columbia University)
Hans B. Dejong (Stanford University)
Robert B. Dunbar (Stanford University)
Dennis A. Hansell (Rosenstiel School of Marine and Atmospheric Science, University of Miami)

Phytoplankton increase projected for the Ross Sea in response to climate change

Posted by mmaheigan

· Thursday, October 26th, 2017

How will phytoplankton respond to climate changes over the next century in the Ross Sea, the most productive coastal waters of Antarctica? Model projections of physical conditions suggest substantial environmental changes in this region, but associated impacts on Ross Sea biology, specifically phytoplankton, remain unclear.

In a recent study, Kaufman et al (2017) generated and analyzed model scenarios for the mid- and late-21^st century using a combination of a biogeochemical model, hydrodynamic simulations forced by a global climate projection, and new data from autonomous gliders. These scenarios indicate increases in the production of phytoplankton in the Ross Sea and increases in the downward flux of carbon in response to environmental changes over the next century. Modeled responses of the different phytoplankton groups to shoaling mixed layer depths shift the biomass composition more towards diatoms by the mid 21^st century. While diatom biomass remains relatively constant in the second half of the 21^st century, the haptophyte Phaeocystis antarctica biomass increases as a result of earlier seasonal sea ice melt, allowing earlier availability of low light, in which P. antarctica thrive.

Modeled climate scenarios for the 21st century project phytoplankton composition changes and increases in primary production and carbon export flux, primarily as a result of shoaling mixed layer depths and earlier available low light.

The projected responses of phytoplankton composition, production, and carbon export to climate-related changes can have broad impacts on food web functioning and nutrient cycling, with wide-ranging potential effects as local deep waters are transported out from the Ross Sea continental shelf. Future changes to this ecosystem have taken on a new relevance as the Ross Sea became home this year to the world’s largest Marine Protected Area, designed to protect critical habitat for highly valued species that rely on the Ross Sea food web. Continued coordination between modeling and autonomous observing efforts is needed to provide vital data for global ocean assessments and to improve our understanding of ecosystem dynamics and climate change impacts in this sensitive and important region.

For other relevant work on observing phytoplankton characteristics in the Ross Sea using gliders, please see: https://doi.org/10.1016/j.dsr.2014.06.011.

And for assimilation of bio-optical glider data in the Ross Sea please see: https://doi.org/10.5194/bg-2017-258.

Authors:
Daniel E. Kaufman (VIMS, College of William and Mary)
Marjorie A. M. Friedrichs (VIMS, College of William and Mary)
Walker O. Smith Jr. (VIMS, College of William and Mary)
Eileen E. Hofmann (CCPO, Old Dominion University)
Michael S. Dinniman (CCPO, Old Dominion University)
John C. P. Hemmings (Wessex Environmental Associates; now at the UK Met Office)

Role for iron in controlling microbial phosphorus acquisition in the ocean

Posted by mmaheigan

· Thursday, October 12th, 2017

In the subtropical North Atlantic, dissolved inorganic phosphorus (DIP) concentrations are depleted and might co-limit N₂ fixation and microbial productivity. There are relatively large pools of dissolved organic phosphorus (DOP), but microbes need an enzyme to access this P source. One such alkaline phosphatase (APase) enzyme requires zinc (Zn) as its activating cofactor. This has been known for almost 30 years. However, recent crystallography studies revealed that two other widespread APase enzymes contain Fe. Via this requirement, Fe availability could regulate microbial access to the DOP pool.

As detailed in a recent publication in Nature Communications (Browning et al. 2017), this hypothesis was tested on a cruise across the tropical North Atlantic by adding Fe and Zn to incubated seawater and monitoring changes in bulk APase using a simple fluorescence assay. Adding Fe significantly increased APase activity in seawater samples collected in areas that were far-removed from coastal and aerosol Fe sources. Despite seawater Zn concentrations being much lower than Fe, it appeared not to be limiting.

Iron (Fe) and zinc (Zn) enrichment experiments conducted in the DIP-depleted tropical North Atlantic suggested that Fe, not Zn, could limit alkaline phosphatase activity (APA). DIP*=DIP–DIN/16, and represents excess DIP availability assuming a 16-fold higher microbial N requirement. Results in the bar chart represent a subset of treatments from one experiment (out of eight conducted).

DIP is depleted in surface waters of the tropical North Atlantic because inputs of North African aerosol Fe stimulates N₂ fixation and leads to microbial drawdown of DIP. If the modern ocean is a good analog for the past, the lack of APase stimulation following experimental Zn addition could reflect limited evolutionary selection for Zn-containing APase. In general, DIP is only substantially depleted where there is enhanced Fe input fueling N₂ fixation; it therefore follows that any significant requirement for APases might be restricted to these relatively high-Fe, low-Zn waters.

On a shorter timescale, growing anthropogenic nitrogen input to the ocean relative to phosphorus could result in more prevalent oceanic phosphorus deficiency. Corresponding iron inputs might then serve as an important control on phosphorus availability for microbes in these regions.

Authors:

Tom Browning (GEOMAR Helmholtz Centre for Ocean Research, Kiel, Germany)
Eric Achterberg (GEOMAR)
Jaw Chuen Yong (GEOMAR)
Insa Rapp (GEOMAR)
Caroline Utermann (GEOMAR)
Anja Engel (GEOMAR)
Mark Moore (Ocean and Earth Science, University of Southampton, Southampton, UK)

Arctic surface waters release methane but also absorb 2,000 times the CO2 for a net cooling effect

Posted by mmaheigan

· Thursday, September 28th, 2017

A recent study by Pohlman et al. published in PNAS showed that ocean waters near the surface of the Arctic Ocean absorbed 2,000 times more carbon dioxide (CO₂) from the atmosphere than the amount of methane released into the atmosphere from the same waters. The study was conducted near Norway’s Svalbard Islands, which overly numerous seafloor methane seeps.

Methane is a more potent greenhouse gas than CO₂, but the removal of CO₂from the atmosphere where the study was conducted more than offset the potential warming effect of the observed methane emissions. During the study, scientists continuously measured the concentrations of methane and CO₂ in near-surface waters and in the air just above the ocean surface. The measurements were taken over methane seeps fields at water depths ranging from 260 to 8530 feet (80 to 2600 meters).

Figure 1. Ocean waters overlying shallow-water methane seeps (white dots) offshore from the Svalbard Islands absorb substantially more atmospheric carbon dioxide than the methane that they emit to the atmosphere. Colors indicate the strength of the negative greenhouse warming potential associated with carbon dioxide influx to these surface waters relative to the positive greenhouse warming potential associated with the methane emissions. Gray shiptracks have background values for the relative greenhouse warming potential.

Analysis of the data confirmed that methane was entering the atmosphere above the shallowest (water depth of 260-295 feet or 80-90 meters) Svalbard margin seeps. The data also showed that significant amounts of CO₂ were being absorbed by the waters near the ocean surface, and that the cooling effect resulting from CO₂ uptake is up to 230 times greater than the warming effect expected from the methane emitted.

Most previous studies have focused only on the sea-air flux of methane overlying seafloor seep sites and have not accounted for the drawdown of CO₂that could offset some of the atmospheric warming potential of the methane. Phytoplankton appeared to be more active in the near-surface waters overlying the seafloor methane seeps, which would explain why so much carbon dioxide was being absorbed. Physical and biogeochemical measurements of near-surface waters overlying the seafloor methane seeps showed strong evidence of upwelling of cold, nutrient-rich waters from depth, stimulating phytoplankton activity and increasing CO₂ drawdown. This study was the first to document this CO₂ drawdown mechanism in a methane source region.

“If what we observed near Svalbard occurs more broadly at similar locations around the world, it could mean that methane seeps have a net cooling effect on climate, not a warming effect as we previously thought,” said USGS biogeochemist John Pohlman, the paper’s lead author. “We are looking forward to testing the hypothesis that shallow-water methane seeps are net greenhouse gas sinks in other locations.”

Authors:
John W. Pohlman (USGS Woods Hole Coastal & Marine Science Center)
Jens Greinert (GEOMAR, Univ. of Tromsø, Royal Netherlands Institute for Sea Research)
Carolyn Ruppel (USGS Woods Hole Coastal & Marine Science Center)
Anna Silyakova (Univ. of Tromsø)
Lisa Vielstädte (GEOMAR)
Michael Casso (USGS Woods Hole Coastal & Marine Science Center)
Jürgen Mienert (Univ. of Tromsø)
Stefan Bünz (Univ. of Tromsø)

Sinking particles as biogeochemical hubs for trace metal cycling and release

Posted by mmaheigan

· Thursday, September 14th, 2017

The extent to which the return of major and minor elements to the dissolved phase in the deep ocean (termed remineralization) is decoupled plays a major role in setting patterns of nutrient limitation in the global ocean. It is well established that major elements such as phosphorus, silicon, and carbon are released at different rates from sinking particles, with major implications for nutrient recycling. Is this also the case for trace metals?

A recent publication by Boyd et al. in Nature Geoscience provides new insights into the biotic and abiotic processes that drive remineralization of metals in the ocean. Particle composition changes rapidly with depth with both physical (disaggregation) and biogeochemical (grazing; desorption) processes leading to a marked decrease in the total surface area of the particle population. The proportion of lithogenic metals in sinking particles also appears to increase with depth, as the biogenic metals may be more labile and hence more readily removed.

Findings from GEOTRACES process studies revealed that release rates for trace elements such as iron, nickel, and zinc vary from each other. Microbes play a key role in determining the turnover rates for nutrients and trace elements. Decoupling of trace metal recycling in the surface ocean and below may result from their preferential removal by microbes to satisfy their nutritional requirements. In addition, the chemistry operating on particle surfaces plays a pivotal role in determining the specific fates of each trace metal. Teasing apart these factors will take time, as there is a complex interplay between chemical and biological processes. Improving our understanding is crucial, as these processes are not currently well represented by state-of-the-art ocean biogeochemical models.

Figure caption: Rapid changes in the characteristics of sinking particles over the upper 200 m as evidenced by: a) differential release of trace metals from sinking diatoms; b) changes in proportion of lithogenic versus biogenic materials; and c) ten-fold decrease in total particle surface area.

Authors:
Philip Boyd (IMAS, Australia)
Michael Ellwood (ANU, Australia)
Alessandro Tagliabue (Liverpool, UK)
Ben Twining (Bigelow, USA)

Relevant links:
GEOTRACES Digest: Iron Superstar

Joint workshop with GEOTRACES in August 2016: Biogeochemical Cycling of Trace Elements within the Ocean

Tiny marine animals strongly influence the carbon cycle

Posted by mmaheigan

· Thursday, August 31st, 2017

What controls the amount of organic carbon entering the deep ocean? In the sunlit layer of the ocean, phytoplankton transform inorganic carbon to organic carbon via a process called photosynthesis. As these particulate forms of organic carbon stick together, they become dense enough to sink out of the sunlit layer, transferring large quantities of organic carbon to the deep ocean and out of contact with the atmosphere.

However, all is not still in the dark ocean. Microbial organisms such as bacteria, and zooplankton consume the sinking, carbon-rich particles and convert the organic carbon back to its original inorganic form. Depending on how deep this occurs, the carbon can be physically mixed back up into the surface layers for exchange with the atmosphere or repeat consumption by phytoplankton. In a recent study published in Biogeosciences, researchers used field data and an ecosystem model in three very different oceanic regions to show that zooplankton are extremely important in determining how much carbon reaches the deep ocean.

Figure 1. Particle export and transfer efficiency to the deep ocean in the Southern Ocean (SO, blue circles), North Atlantic Porcupine Abyssal Plain site (PAP, red squares) and the Equatorial Tropical North Pacific (ETNP, orange triangles) oxygen minimum zone. a) particle export efficiency of fast sinking particles (Fast PEeff) against primary production on a Log10 scale. b) transfer efficiency of particles to the deep ocean expressed as Martin’s b (high b = low efficiency). Error bars in b) are standard error of the mean for observed particles, error too small in model to be seen on this plot.

In the Southern Ocean (SO), zooplankton graze on phytoplankton and produce rapidly sinking fecal pellets, resulting in an inverse relationship between particle export and primary production (Fig. 1a). In the North Atlantic (NA), the efficiency with which particles are transferred to the deep ocean is comparable to that of the Southern Ocean, suggesting similar processes apply; but in both regions, there is a large discrepancy between the field data and the ecosystem model (Fig. 1b), which poorly represents particle processing by zooplankton. Conversely, much better data-model matches are observed in the equatorial Pacific, where lower oxygen concentrations mean fewer zooplankton; this reduces the potential for zooplankton-particle interactions that reduce particle size and density, resulting in a lower transfer efficiency.

This result suggests that mismatches between the data and model in the SO and NA may be due to the lack of zooplankton-particle parameterizations in the model, highlighting the potential importance of zooplankton in regulating carbon export and storage in the deep ocean. Zooplankton parameterizations in ecosystem models must be enhanced by including zooplankton fragmentation of particles as well as consumption. Large field programs such as EXPORTS could help constrain these parameterisation by collecting data on zooplankton-particle interaction rates. This will improve our model estimates of carbon export and our ability to predict future changes in the biological carbon pump. This is especially important in the face of climate-driven changes in zooplankton populations (e.g. oxygen minimum zone (OMZ) expansion) and associated implications for ocean carbon storage and atmospheric carbon dioxide levels.

Authors:
Emma L. Cavan (University of Tasmania)
Stephanie A. Henson (National Oceanography Centre, Southampton)
Anna Belcher (University of Southampton)
Richard Sanders (National Oceanography Centre, Southampton)

Phytoplankton can actively diversify their migration strategy in response to turbulent cues

Posted by mmaheigan

· Thursday, August 17th, 2017

Turbulence is known to be a primary determinant of plankton fitness and succession. However, open questions remain about whether phytoplankton can actively respond to turbulence and, if so, how rapidly they can adapt to it. Recent experiments have revealed that phytoplankton can behaviorally respond to turbulent cues with a rapid change in shape, and this response occurs over a few minutes. This challenges a fundamental paradigm in oceanography that phytoplankton are passively at the mercy of turbulence.

Phytoplankton are photosynthetic microorganisms that form the base of most aquatic food webs, impact global biogeochemical cycles, and produce half of the world’s oxygen. Many species of phytoplankton are motile and migrate in response to gravity and light levels: Upward toward light during the day to photosynthesize and downward at night toward higher nutrient concentrations. Disruption of this diurnal migratory strategy is an important contributor to the succession between motile and non-motile species when conditions become more turbulent. However, this classical view neglects the possibility that motile species can actively respond in an effort to avoid layers of strong turbulence. A recent study by Sengupta, Carrara and Stocker, published in Nature has shown that some raphidophyte and dinoflagellate phytoplankton can actively diversify their migratory strategy in response to hydrodynamic cues characteristic of overturning by the smallest turbulent eddies in the ocean. Laboratory experiments in which cells experienced repeated overturning with timescales and statistics representative of ocean turbulence revealed that over timescales as short as ten minutes, an upward-swimming population split into two subpopulations, one swimming upward and one swimming downward. Quantitative morphological analysis of the harmful algal bloom-forming raphidophyte Heterosigma akashiwo revealed that this behavior was accompanied by a change in cell shape, wherein the cells that changed their swimming direction did so by going from an asymmetric pear shape to a more symmetric egg shape. A model of cell mechanics showed that the magnitude of this shift was minute, yet sufficient to invert the cells’ preferential swimming direction. The results highlight the advanced level of control that phytoplankton have on their migratory behavior.

Understanding how fluctuations in the oceans’ turbulence landscape impacts phytoplankton is of fundamental importance, especially for predicting species succession and community structure given projected climate-driven changes in temperature, winds, and upper ocean structure.

An upward-swimming phytoplankton population splits into upward- and downward-swimming sub-populations when exposed to turbulent eddies, due to a subtle change in cell shape. Illustration by: A. Sengupta, G. Gorick, F. Carrara and R. Stocker

This work was co-funded by a Human Frontier Science Program Cross Disciplinary Fellowship (LT000993/2014-C to A.S.), a Swiss National Science Foundation Early Postdoc Mobility Fellowship (to F.C.), and a Gordon and Betty Moore Marine Microbial Initiative Investigator Award (GBMF 3783 to R.S.)

Untangling the mystery of domoic acid events: A climate-scale perspective

Posted by mmaheigan

· Thursday, August 3rd, 2017

The diatom Pseudo-nitzchia produces a neurotoxin called domoic acid, which in high concentrations affects wildlife ranging from mussels and crabs to seabirds and sea lions, as well as humans. In humans, the effects of domoic acid poisoning can range from gastrointestinal distress to memory loss, and even death. Despite being studied in laboratories since the late 1980s, there is no consensus on the environmental conditions that lead to domoic acid events. These events are most frequent and impactful in eastern boundary current regions such as the California Current System, which is bordered by Washington, Oregon, and California. In Oregon alone, there have been six major domoic acid events: 1996, 1998-1999, 2001, 2002-2006, 2010, 2014-2015. McKibben et al. (2017) investigated the regulation of domoic acid at a climate scale to develop and test an applied risk model for the US West Coast” to read “McKibben et al. (2017) investigated the regulation of domoic acid at regional and decadal scales in order to develop and test an applied risk model for the impact of climate on the US West Coast. They used the PDO and ONI climate variability indices, averages of monthly and 3 month running means of SST anomaly values and variability to look at basin-scale ocean conditions. At a local scale, data were from zooplankton sampling every two to four weeks between 1996 to 2015 at hydrographic station offshore of Newport, OR. Additionally, the NOAA NCDC product “Daily Optimum Interpolation, Advanced Very High Resolution Radiometer Only, Version 2, Final+Preliminary SST” was used to obtain the monthly SST anomaly metric, based on combined in situ and satellite data.

(A) Warm and cool ocean regimes, (B) local SST anomaly, and (C and D) biological response. (A) PDO (red or blue vertical bars) and ONI (black line) indices; strong (S) to moderate (M) El Nino (+1) and La Nina (−1) events are labeled. (B) SST anomaly 20 nm off central OR. (C) The CSR anomaly 5 nm off central OR. (D) Monthly OR coastal maximum DA levels in razor clams (vertical bars); horizontal black line is the 20-ppm closure threshold. Black line in D shows the spring biological transition date (right y axis). At the top of the figure, black boxes indicate the duration of upwelling season each year; red vertical bars indicate the timing of annual DA maxima in relationship to upwelling. Gray shaded regions are warm regimes based on the PDO. Dashed vertical lines indicate onset of the six major DA events. The September 2014 arrival of the NE Pacific Warm Anomaly (colloquially termed “The Blob”) to the OR coastal region is labeled on B. “X” symbols along the x axes indicate that no data were available for that month (B–D).

Their findings show that these events have occurred when there is advection of warmer water masses onto the continental shelf from southern or offshore areas. When the warm phase of the Pacific Decadal Oscillation (PDO) and El Niño coincide, the effect is additive. In the warm regime years, there is a later spring biological transition date, weaker alongshore currents, elevated water temperatures, and plankton communities are dominated by subtropical rather than subarctic species. The authors also note relative differences between the prevalence and phenology of domoic acid events in OR, CA and WA, which warrants further study via regional-scale modeling. Overall, this research shows a clear and enhanced risk of toxicity in shellfish during warm phases of natural climate oscillations. If predictions of more extreme warming come to bear, this would potentially lead to increased DA event intensity and frequency in coastal zones around the globe. This will not only affect wildlife, but may cause significant closures of economically important fisheries (e.g., Dungeness crab, anchovy, mussel, and razor clam), which would impact local communities and native populations.

Authors:
Morgaine McKibben (Oregon State Univ., NOAA Northwest Fisheries Science Center)
William Peterson (NOAA Northwest Fisheries Science Center)
Michelle Wood (Univ. Oregon)
Vera L. Trainer (NOAA Northwest Fisheries Science Center)
Matthew Hunter (Oregon Dept. Fish & Wildlife)
Angelicque E. White (Oregon State Univ.)

An autonomous approach to monitoring coral reef health

Posted by mmaheigan

· Thursday, July 20th, 2017

Coral reefs are diverse, productive ecosystems that are highly vulnerable to changing ocean conditions such as acidification and warming. Coral reef metabolism—in particular the fundamental ecosystem properties of net community production (NCP; the balance of photosynthesis and respiration) and net community calcification (NCC; the balance of calcification and dissolution)—has been proposed as a proxy for reef health. NCC is of particular interest, since ocean acidification is expected to have detrimental effects on reef calcification.

Traditionally, these metabolic rates are quantified through laborious methods that involve discrete sampling, which, due to a limited number of observations, often fails to characterize natural variability on time scales of minutes to days. In a recent paper in JGR, Takeshita et al. (2016) presented the Benthic Ecosystem and Acidification Measurement System (BEAMS), a fully autonomous system that simultaneously measures NCP and NCC at 15-minute intervals over a period of weeks. BEAMS utilizes the gradient flux method to quantify benthic metabolic rates by measuring chemical (pH and O₂) and velocity gradients in the turbulent benthic boundary layer.

Two BEAMS were simultaneously deployed on Palmyra Atoll located approximately one km apart over vastly different benthic communities. One site was a healthy reef with approximately 70% coral cover, and the other was a degraded reef site with only 5% coral cover that was dominated by a non-calcifying invasive corallimorph Rhodactis howesii. Over the course of two weeks, BEAMS collected over 1,000 measurements of NCP and NCC from each site, yielding significantly different ratios of NCP to NCC between the two sites. These initial results suggest that BEAMS is capable of detecting different metabolic states, as well as patterns consistent with degrading reef health.

BEAMS is an exciting new autonomous tool to monitor reef health and study drivers of reef metabolism on timescales ranging from minutes to months (and potentially years). Additionally, autonomous measurement tools increase the potential for widespread and comparable observations across reefs and reef systems. Such knowledge will greatly improve our ability to predict the fate of coral reefs in a changing ocean.

Authors:
Yui Takeshita (Monterey Bay Aquarium Research Institute)

Funding for the Ocean Carbon & Biogeochemistry Project Office is provided by the National Science Foundation (NSF) and the National Aeronautics and Space Administration (NASA). The OCB Project Office is housed at the Woods Hole Oceanographic Institution.

Archive for New OCB Research – Page 24

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