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Archive for New OCB Research – Page 24

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 CO2 from 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 CO2 that 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)

The changing ocean carbon cycle

Posted by mmaheigan 
· Thursday, July 6th, 2017 

Since preindustrial times, the ocean has removed from the atmosphere 41% of the carbon emitted by human industrial activities (Figure 1). The globally integrated rate of ocean carbon uptake is increasing in response to rising atmospheric CO2 levels and is expected to continue this trend for the foreseeable future. However, the inherent uncertainties in ocean surface and interior data associated with ocean carbon uptake processes make it difficult to predict future changes in the ocean carbon sink. In a recent paper, McKinley et al. (2017), review the mechanisms of ocean carbon uptake and its spatiotemporal variability in recent decades. Looking forward, the potential for direct detection of change in the ocean carbon sink, as distinct from interannual variability, is assessed using a climate model large ensemble, a novel approach to studying climate processes with an earth systems model, the “large ensemble.” In a large ensemble, many runs of the same model are done so as to directly distinguish natural variability from long-term trends.


This analysis illustrates that variability in CO2 flux is large enough to prevent detection of anthropogenic trends in ocean carbon uptake on at least decadal to multi-decadal timescales, depending on location. Earliest detection of trends is most attainable in regions where trends are expected to be largest, such as the Southern Ocean and parts of the North Atlantic and North Pacific. Detection will require sustained observations over many decades, underscoring the importance of traditional ship-based approaches and integration of new autonomous observing platforms as part of a global ocean carbon observing system.

Please see a relevant OCB outreach tool on ocean carbon uptake developed by McKinley and colleagues:
OCB teaching/outreach slide deck Temporal and Spatial Perspectives on the Fate of Anthropogenic Carbon: A Carbon Cycle Slide Deck for Broad Audiences  – also download explanatory notes

Quantifying coastal and marine ecosystem carbon storage potential for climate mitigation policy and management

Posted by mmaheigan 
· Wednesday, June 21st, 2017 

Under the increasing threat of climate change, conservation practitioners and policy makers are seeking innovative and data–driven recommendations for mitigating emissions and increasing natural carbon sinks through nature-based solutions. While the ocean and terrestrial forests, and more recently, coastal wetlands, are well known carbon sinks, there is interest in exploring the carbon storage potential of other coastal and marine ecosystems such as coral reefs, kelp forests, phytoplankton, planktonic calcifiers, krill, and teleost fish. A recent study in Frontiers in Ecology and the Environment reviewed the potential and feasibility of managing these other coastal and marine ecosystems for climate mitigation. The authors concluded, that while important parts of the carbon cycle, coral reefs, kelp forests, planktonic calcifiers, krill, and teleost fish do not represent long-term carbon stores, and in the case of fish, do not represent a sequestration pathway. Phytoplankton do sequester globally significant amounts of carbon and contribute to long-term carbon storage in the deep ocean, but there is currently no good way to manage them to increase their carbon storage capacity; additionally, the vast majority of phytoplankton is located in international waters that are outside national jurisdictions, making it very difficult to include them in current climate mitigation policy frameworks.

Comparatively, coastal wetlands (mangroves, tidal marshes, and seagrasses) effectively sequester carbon long-term (up to 10x more carbon stored per unit area than terrestrial forests with 50-90% of the stored carbon residing in the soil), and fall within clear national jurisdictions, which facilitates effective and quantifiable management actions. In addition, wetland degradation has the potential to release vast amounts of stored carbon back into the atmosphere and water column, meaning that conservation and restoration of these systems can also reduce potential emissions. The authors conclude that coastal wetland protection and restoration should be a primary focus in comprehensive climate change mitigation plans along with reducing emissions.

Authors:
Jennifer Howard (Conservation International)
Ariana Sutton-Grier (University of Maryland, NOAA)
Dorothée Herr (IUCN)
Joan Kleypas (NCAR)
Emily Landis (The Nature Conservancy)
Elizabeth Mcleod (The Nature Conservancy)
Emily Pidgeon (Conservation International)
Stefanie Simpson (Restore America’s Estuaries)

Original paper: http://onlinelibrary.wiley.com/doi/10.1002/fee.1451/full

Winter ventilation depth constrains the impact of the biological pump on CO2 uptake in the North Pacific Ocean

Posted by mmaheigan 
· Thursday, June 8th, 2017 

The North Pacific accounts for ~25% of the global ocean’s uptake of carbon dioxide (CO2) from the atmosphere. However, the relative importance of the biological pump vs. physical circulation in driving ocean uptake of CO2 remains poorly understood.

In a recent study, Palevsky and Quay (2017) used geochemical measurements collected on sixteen container ship transects between Hong Kong and Long Beach, CA to evaluate the drivers of CO2 uptake across 8,000 kilometers in the North Pacific basin over the full annual cycle. In the eastern North Pacific, biologically-driven export of organic carbon below the winter ventilation depth fully offsets the uptake of CO2 from the atmosphere. However, in the Kuroshio region of the western North Pacific, which has a deep winter mixed layer, the majority of the organic carbon exported during the productive summer season is subsequently respired and ventilated back to the atmosphere in winter. Subsequently, biologically-driven export offsets only a small fraction of the CO2 uptake by the ocean and, instead, physical transport is the dominant process removing inorganic carbon from the region.

We further show that that mechanistic coupling between biological carbon export and ocean uptake of CO2 from the atmosphere is sensitive to the seasonal timing of biological export and ventilation, as well as the magnitude of export. Future studies therefore need to measure biological carbon export and ventilation throughout the full annual cycle in order to better understand controls on regional variations in ocean CO2 uptake rates and future changes in these rates.

Data from 16 shipboard transects across the North Pacific revealed a basin-wide gradient between the Kuroshio and Eastern regions in the relative roles of biological vs. physical processes in removing dissolved inorganic carbon from the surface ocean.

 

Authors:
Hilary I. Palevsky (Woods Hole Oceanographic Institution)
Paul D. Quay (University of Washington)

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shorelines siderophore silica silicate silicon cycle sinking sinking particles size SOCCOM soil carbon southern ocean south pacific spatial covariations speciation SST state estimation stoichiometry subduction submesoscale subpolar subtropical sulfate surf surface surface ocean Synechococcus technology teleconnections temperate temperature temporal covariations thermocline thermodynamics thermohaline thorium tidal time-series time of emergence titration top predators total alkalinity trace elements trace metals trait-based transfer efficiency transient features trawling Tris trophic transfer tropical turbulence twilight zone upper ocean upper water column upwelling US CLIVAR validation velocity gradient ventilation vertical flux vertical migration vertical transport warming water clarity water mass water quality waves weathering western boundary currents wetlands winter mixing zooplankton

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