Archive for plankton

Mixotrophs in the northern North Atlantic

Posted by mmaheigan 
· Tuesday, April 16th, 2024 

Mixotrophs (or mixoplankton) are now accepted as a third group of plankton alongside phytoplankton and zooplankton. Our knowledge of mixotrophs lags far behind that of the other two groups. We currently have only a limited understanding of mixotrophs’ biogeographical distribution across ocean basins, and what environmental factors are associated with their distribution.

The authors of a study recently published in Frontiers in Marine Science reviewed nearly 230,000 individual microplankton samples collected by the North Atlantic Continuous Plankton Recorder program between 1958 and 2015 and calculated the proportion of organisms that are considered mixotrophs in each sample. They classified protist species in the dataset as phytoplankton, mixotrophs, or microzooplankton (heterotrophs), based on existing literature. Taken together across seasonsin shelf waters (depth ≤ 300m), mixotrophs comprise a greater proportion of the microplankton community when nitrate is high and photosynthetically available radiation (PAR) is low (e.g. during the late fall and winter), or when nitrate is low and PAR is moderate to high (e.g. during the summer and early fall). When both nitrate and PAR are high, mixotrophs comprise less of the community compared to phytoplankton. The same pattern was found in offshore waters (depth > 300m), but the key macronutrient was phosphate rather than nitrate. The annual average proportion of mixotrophs in microplankton samples compared to phytoplankton has increased since 1958 in the offshore portion of the study region, with a notable changepoint in 1993; this increasing trend is strongest in the winter season.

This paper is useful for aquatic ecologists who want to integrate mixotrophic plankton into their understanding of marine food webs and biogeochemical cycles. Understanding mixotroph temporal and spatial distributions, as well as the environmental conditions under which they flourish, is imperative to understanding their impact on trophic transfer and biogeochemical cycling.

Authors
Karen Stamieszkin (Bigelow Laboratory for Ocean Sciences)
Nicole Millette (Virginia Institute of Marine Science)
Jessica Luo (NOAA Geophysical Fluid Dynamics Laboratory)
Elizabeth Follett (University of Liverpool)
Nick Record (Bigelow Laboratory of Ocean Science)
David Johns (Marine Biological Association)

 

Backstory
This work and the collaboration that made it possible was catalyzed by the Eco-DAS XII symposium, attended by Karen Stamieszkin, Nicole Millette, Jessica Luo, and Elizabeth Follett in 2016. Nicole had an idea for an analysis but lacked collaborators, just as she was ready to give up on it, Karen, Jessica, and Elizabeth expressed interest in the project. Karen, Jessica, and Elizabeth each brought a unique perspective that helped make Nicole’s original idea more practical and ensured that the analysis would come to life.

The collaboration that began with this paper lead to the OCB Mixotrophs & Mixotrophy Working Group led by Karen, Jessica, and Nicole, and a successful grant proposal to study mixotrophy awarded to Nicole and Karen by NSF’s Biological Oceanography program. This story shows the importance and power of programs that connect researchers across disciplines, especially in the early stages of their careers.

Ocean Acidification drives shifts in global stoichiometry and carbon export efficiency

Posted by mmaheigan 
· Friday, November 19th, 2021 

Marine food webs and biogeochemical cycles react sensitively to increases in carbon dioxide (CO2) and associated ocean acidification, but the effects are far more complex than previously thought. A comprehensive study published in Nature Climate Change by a team of researchers from GEOMAR dove deep into the impacts of ocean acidification on marine biota and biogeochemical cycling. The authors combined data from five large-scale field experiments with natural plankton communities to investigate how carbon cycling and export respond to ocean acidification.

The biological pump is a key mechanism in transferring carbon to the deep ocean and contributes significantly to the oceans’ function as a carbon sink. The carbon-to-nitrogen ratio of sinking biogenic particles, here termed (C:Nexport), determines the amount of carbon that is transported from the euphotic zone to the ocean interior per unit nutrient, thereby controlling the efficiency of the biological pump. The authors demonstrate for the first time that ocean acidification can change the elemental composition of organic matter export, thereby potentially altering the biological pump and carbon sequestration in a future ocean (Figure 1).

Figure 1: Until now, the common assumption is that changes in C:N (and biogeochemistry, in general) are mainly driven by phytoplankton. In a series of in situ mesocosm experiments in different biomes (left), Taucher et al., (2020) found distinct impacts of ocean acidification on the C:N ratio of sinking organic matter (middle). By linking these observations to analysis of plankton community composition, the authors found a key role of heterotrophic processes in controlling the response of C:N to OA, particularly by altering the quality and carbon content of sinking organic matter within the biological pump (right).

Surprisingly, the observed responses were highly variable: C:Nexport increased or decreased significantly with increasing CO2, depending on the composition of species and the structure of the food web. The authors found that heterotrophic processes driven by bacteria and zooplankton play a key role in controlling the response of C:Nexport to ocean acidification. This contradicts the widespread paradigm that primary producers are the principal driver of biogeochemical responses to ocean change.

Considering that such diverse pathways, by which planktonic food webs shape the elemental composition and biogeochemical cycling of organic matter, are not represented in state-of-the-art earth system models, these findings also raise the question: Are currently able to predict the large-scale consequences of ocean acidification with any certainty?

 

Authors:
Jan Taucher (GEOMAR, Kiel, Germany)
Tim Boxhammer (GEOMAR, Kiel, Germany)
Lennart T. Bach (University of Tasmania, Hobart, Australia)
Allanah J. Paul (GEOMAR, Kiel, Germany)
Markus Schartau (GEOMAR, Kiel, Germany)
Paul Stange (GEOMAR, Kiel, Germany)
Ulf Riebesell (GEOMAR, Kiel, Germany)

The role of nutrient trapping in promoting shelf hypoxia in the southern Benguela upwelling system

Posted by mmaheigan 
· Thursday, September 3rd, 2020 

The southern Benguela upwelling system (SBUS) off southwest Africa is an exceptionally fertile ocean region that supports valuable commercial fisheries. The productivity of this system derives from the upwelling of nutrient-rich Subantarctic Mode Water, and from the concurrent entrainment of nutrients regenerated proximately on the expansive continental shelf. The SBUS is prone to severe seasonal hypoxic events that decimate regional fisheries, occurrences of which are inextricably linked to the inherent nutrient dynamics. In a study recently published in JGR Oceans, the authors sought to understand the mechanisms sustaining elevated concentrations and seasonally-variable distributions of nutrients in the SBUS, in relation to the subsurface oxygen content. Inter-seasonal measurements of nutrients and nitrate isotope ratios across the SBUS in 2017 revealed that upwards of 48% (summer) and 63% (winter) of the on‐shelf nutrients derived from regeneration in situ.  The severity of hypoxia at the shelf bottom, in turn, correlated with the incidence of regenerated nutrients. The accrual of nutrients at the shelf bottom appears to be aided by hydrographic fronts that restrict offshore transport, trapping regenerated nutrients on the SBUS shelf and increasing the pool of nutrients available for upwelling – ultimately contributing to hypoxic events. This study underscores the need – if we are to develop a mechanistic and predictive understanding of hypoxia in the SBUS and elsewhere – to elucidate the role of shelf circulation in promoting the accrual of regenerated nutrients on the continental shelf. The next step is to combine new and existing observations with quantitative simulations to further interrogate the coupled physical-biogeochemical mechanisms that modulate the intensity of hypoxia.

Figure caption: Schematic of proposed nutrient-trapping mechanism: Deep nutrient-rich Subantarctic Mode Water (SAMW) acquires more nutrients as it passes over the shelf sediments from the regeneration of exported particulate organic material (POM). The production of this POM is fueled by nutrients stripped from the surface waters advecting back off-shore. The thickness of the arrows represents nutrient concentrations. Triangles indicate the positions of the Shelf Break Front (SBF) and Columbine Front (CF), coincident with an observed subduction of the Ekman layer and downwelling at the inner front boundary.

Authors
Raquel Flynn (University of Cape Town)
Julie Granger (University of Connecticut)
Jennifer Veitch (South African Environmental Observation Network)
Samantha Siedlecki (University of Connecticut)
Jessica Burger (University of Cape Town)
Keshnee Pillay (South Africa Department of Environment, Forestry and Fisheries)
Sarah Fawcett (University of Cape Town)

Marine Snowfall at the Equator

Posted by mmaheigan 
· Thursday, July 19th, 2018 

The continual flow of organic particles such as dead organisms and fecal material towards the deep sea is called “marine snow,” and it plays an important role in the ocean carbon cycle and climate-related processes. This snowfall is most intense where high primary production can be observed near the surface. This is the case along the equator in the Pacific and Atlantic Oceans. However, it is not well known how particles are distributed at depth and which processes influence this distribution. A recent study published in Nature Geoscience involved the use of high-resolution particle density data using the Underwater Vision Profiler (UVP) from the equatorial Atlantic and Pacific Oceans down to a depth of 5,000 meters, revealing that several previously accepted ideas on the downward flux of particles into the deep sea should be revisited.

Figure 1. The Underwater Vision Profiler (UVP) during a trial in the Kiel Fjord. The UVP provided crucial data for the new study. Photo: Rainer Kiko, GEOMAR

 

It is typically assumed that the largest particle density can be found close to the surface and that density attenuates continuously with depth. However, high-resolution particle data show that density increases again in the 300-600-meter depth range. The authors attribute this observation to the daily migratory behavior of organisms such as zooplankton that retreat to these depths during the day, contributing to the particle load via defecation and mortality.

Another surprising result is the observation of many small particles below 1,000 meters depth that contribute a large fraction of the bathypelagic particle flux. This observation counters the general assumption, especially in many biogeochemical models, that particle flux at depth comprises fast sinking particles such as fecal pellets. Diminished remineralization rates of small particles or increased disaggregation of larger particles may contribute to the elevated small particle fluxes at this depth.

Figure 2. Zonal current velocity and Particulate Organic Carbon (POC) content across the equatorial Atlantic at 23˚W as observed in November 2012. From left to right: Zonal current velocity, POC content in small particle fraction and POC content in large particle fraction (adapted from Kiko et al. 2017).

 

This study highlights the importance of coupled biological and physical processes in understanding and quantifying the biological carbon pump. Further work on this important topic can now also be submitted to the new Frontiers in Marine Science research topic “Zooplankton and Nekton: Gatekeepers of the Biological Pump” (https://www.frontiersin.org/research-topics/8114/zooplankton-and-nekton-gatekeepers-of-the-biological-pump; Co-editors R. Kiko, M. Iversen, A. Maas, H. Hauss and D. Bianchi). The research topic welcomes a broad range of contributions, from individual-based process studies, to local and global field observations, to modeling approaches to better characterize the role of zooplankton and nekton for the biological pump.

 

Authors:
R. Kiko (GEOMAR)
A. Biastoch (GEOMAR)
P. Brandt (GEOMAR, University of Kiel)
S. Cravatte (LEGOS, University of Toulouse)
H. Hauss (GEOMAR)
R. Hummels (GEOMAR)
I. Kriest (GEOMAR)
F. Marin (LEGOS, University of Toulouse)
A. M. P. McDonnell (University of Alaska Fairbanks)
A. Oschlies (GEOMAR)
M. Picheral (Laboratoire d’Océanographie de Villefranche-sur-Mer, Observatoire Océanologique)
F. U. Schwarzkopf (GEOMAR)
A. M. Thurnherr (Lamont-Doherty Earth Observatory,)
L. Stemmann (Sorbonne Universités, Observatoire Océanologique)

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.)

Trace metal uptake and remineralization and their impact on upper ocean stoichiometry

Posted by mmaheigan 
· Wednesday, July 6th, 2016 

1. Stoichiometry of metals in the ocean

The close relationship between the stoichiometry of nutrients dissolved in the upper ocean and the planktonic organisms that grow in these waters has long been recognized (1, 2). The stoichiometry of 106 C:16 N:1 P first summarized by Redfield has become a fundamental concept of marine biogeochemistry, with numerous studies using the ratio as a benchmark to assess ecosystem function. Decades after the work of Redfield, with the implementation of trace metal-clean techniques, oceanographers produced the first meaningful measurements of dissolved trace metals in the open ocean (3-5), and they found that many of the bioactive metals such as Fe, Zn, Ni, Cu and Cd are also depleted in surface waters and enriched at depth, similar to the macronutrients. Such nutrient-like behavior supported not only a growing understanding of the physiological roles that these metals play in phytoplankton physiology (6), but it also indicated that biological uptake and sub-surface remineralization were important processes for controlling the distributions of these bioactive metals in the ocean. Thus, the biogeochemical behavior of the micronutrient metals is in many ways analogous to that of the macronutrients N, P and Si.

In the open ocean far from coastal and shelf influences, dissolved concentrations of bioactive metals increase with depth at relatively consistent ratios to macronutrients (5), and these metal:nutrient remineralization ratios have been used to approximate the composition of sinking biogenic material and euphotic zone phytoplankton (7, 8). These ‘extended Redfield ratios’ have been compared to average compositions of marine phytoplankton species grown in culture (9-12),  and the general agreement between these approaches further supports the importance of biological uptake and subsequent remineralization of trace metals in the upper ocean as key processes impacting trace metal geochemistry. Average metal:nutrient stoichiometries for phytoplankton have also been compared to dissolved stoichiometries in the ambient water, and relationships between these fractions have been used to estimate nutrient limitation and deficiency in the ocean (13). Thus, there is significant interest in controls on upper ocean metal stoichiometries, as well as the relationships between cellular/biological, particulate and dissolved fractions.

Analogous to macronutrients, there are also relationships between metal stoichiometries in phytoplankton and those in deeper waters of the ocean. Departures from these relationships are likely to provide insights into the internal biogeochemical cycling of metals in the ocean. Morel and Hudson (7) noted differences in the extended stoichiometries of plankton and the water column and concluded that they must reflect the relative efficiency of remineralization of the elements, as well as the propensity of elements to be scavenged onto sinking particles in the sub-surface ocean. Similarly, the rapid remineralization of trace metals from sinking plankton was addressed in seminal work by Collier and Edmond (14). Using carefully collected data on surface plankton material, and with more computational rigor than (7), they compared surface particle stoichiometries to deep water dissolved stoichiometries and calculated the relative remineralization of plankton-associated elements in sinking biogenic material. They noted significant differences among the behaviors of biogenic metals such as Cd, Ni and Fe due to their scavenging and remineralization behaviors. More recently, Morel (15) mused about these processes and their relationships to cellular biochemistry and evolution of phytoplankton physiology and ocean biogeochemistry.

Through the GEOTRACES program, the data to test and extend these early, relatively simple box models and stoichiometric comparisons are now available. Metal concentrations and stoichiometries for phytoplankton, bulk and size-fractionated particulate material, and co-located dissolved species have been measured in the North Atlantic and South Pacific Oceans thus far. Combined with data for non-bioactive metals such as Ti and Th, these data also provide the opportunity to discern the behavior and contributions of lithogenic vs. biogenic matter, as well as the processes of remineralization and scavenging.

2. Processes affecting dissolved and particulate stoichiometries of trace metals

Vertical profiles of dissolved macronutrients show characteristic depletion at the surface and enrichment at depth due to remineralization, and dissolved micronutrients often show the same behavior. However, the internal cycling of metals in the ocean is expected to differ from that of macronutrients for a few salient reasons. Some metals such as Fe are significantly less soluble than macronutrients and are prone to abiotic adsorption onto particulate surfaces (16). This process is driven by thermodynamics, and the accompanying process of desorption also occurs; the net observed process is typically called ‘scavenging’ (Fig. 1). Scavenging in the deep ocean causes concentrations of less soluble metals such as Fe and Al to decrease along the path of thermohaline circulation, in contrast to macronutrients and more soluble metals that may mimic macronutrient behavior such as Cd and Zn (17). In the absence of significant lateral nutrient inputs, the balance of scavenging and remineralization will influence the resulting vertical profiles of dissolved elements (18).

Another key difference between macronutrients and metals is the importance of abiotic particulate fractions such as lithogenic (e.g., aeolian dust and sediment) and authigenic (e.g., Fe- and Mn-oxyhydroxide) phases. While biogenic phases are almost universally produced at the surface and remineralized with depth, abiotic phases can exhibit very different and dynamic internal cycles (19). Dust events, lateral transport and poorly constrained scavenging processes can both deliver and remove specific metals alongside biological processes. Lithogenic phases are generally denser and more refractory than biogenic particles and detritus and are thought to sink more rapidly and remineralize more slowly and at greater depth (Fig. 1; 20). Lithogenic particles may also (re)scavenge metals differently than biogenic material. Efforts to examine these processes in sinking material have been extremely limited to date, with only a few studies examining metals in trace metal-clean sediment traps (21, 22). However, recently published datasets from the GEOTRACES program are shedding new light on the multiple facets of metal partitioning and how they affect subsurface remineralization and scavenging.

A comparison of metal:phosphorus ratios in the upper ocean illuminates some of these processes. Figure 2 displays Cd:P, Fe:P, Co:P and Ni:P ratios in particles in the upper 100m, 100-300m, and 300-1,000m of the water column in the middle of the North Atlantic basin. Particulate material is sub-divided into ratios for phytoplankton cells and non-lithogenic particles (corrected for lithogenic minerals using Ti; 19). Also plotted are dissolved remineralization ratios (that is, the slope of a linear regression between the dissolved metal and phosphate) for these upper ocean depth ranges. The close coupling of Cd and P biogeochemistry has long been recognized (4), and indeed we observe very close agreement (within a factor of about 2) between dissolved Cd:P remineralization and Cd:P in surface ocean particles, as well subsurface particles. Clearly these elements are remineralizing from sinking particles at similar rates. Such comparisons of particulate and dissolved constituents need to carefully consider the different residence times of these fractions and the likelihood for lateral inputs. Here, we have chosen to focus on stations from the mid-North Atlantic gyre, where the upper 700m of the water column consists primarily of a single water mass (23).

In contrast, the remineralization of Fe and P are quickly decoupled in the water column (Fig. 2). Between 100 and 300m, typically the depth of most rapid regeneration of sinking organic material, labile particulate Fe:P has more than doubled from that in surface waters, and the Fe:P ratio of remineralized dissolved elements (0.98 mmol/mol) is more than 10-fold below that of the labile material that is sinking into these waters. Looking deeper into the water column, Fe and P continue to decouple in labile (i.e., non-lithogenic) particulates, with Fe:P of 300-1,000m particles increasing 10-fold and the dissolved remineralization ratio being nearly 1,000-fold lower (0.35 mmol/mol). Additionally, organic ligands play an important role in stabilizing dissolved Fe (24), so dissolved Fe and P ratios may be further decoupled by biological processes impacting the production and fate of these ligands (20).

A strength of GEOTRACES datasets is their wide coverage of the periodic table, and additional insights can be gained from looking at the behaviors of other bioactive trace metals that are also incorporated into sinking biogenic material. Co:P ratios in particles and remineralized dissolved fractions in the water column follow the same trend as Fe, but the decoupling of Co and P is much more subtle than with Fe, presumably due to differences in ligand coordination and Co co-oxidation with Mn (25, 26). Dissolved Co:P remineralization ratios at 100-300m generally match those found in phytoplankton and drop only 3-fold below 300m. Similarly, labile particulate Co:P ratios don’t change between 0-100m and 100-300m, also indicating that Co and P remineralize in tandem in the upper 300m. Below 300m, labile particulate Co:P increases approximately 3-fold (in contrast with Fe:P, which increases 12-fold), and this depth effect matches the effect in dissolved remineralization ratios. Thus, even though Fe and Co are considered hybrid metals that display both biological uptake and scavenging, there are clear differences in the behaviors of these metals. Nickel provides yet another perspective on the coupling of metals and P. Dissolved remineralization ratios in both subsurface depth ranges closely resemble surface ocean labile particles, supporting the biological coupling of Ni and P (5). However, residual labile particulate Ni:P increases 2- to 4-fold in successive depth ranges, indicating that remineralization is rather decoupled. Given that Ni seems to be associated with both organic material and opal frustules in diatoms (27), it may be that Ni and P are remineralized from particulate organic matter in tandem, but some Ni remains associated with sinking biogenic silica in the ocean.

3. Additional tools to explore and differentiate remineralization processes

The GEOTRACES program has welcomed the application of new analytical approaches that further enable us to study the cycling of metals in the ocean. Spectroscopy and quantitative imaging methods using synchrotron radiation have become more common in the past decade (28), and these allow us to analytically distinguish the behaviors of different fractions of particle assemblages. During the FeCycle II project, a GEOTRACES process study, the fate of Fe was tracked during a spring diatom bloom (29). Diatom cells from the dominant bloom species (Asterionellopsis glacialis) were collected in surface waters and from trace-metal clean sediment traps at 100m and 200m in the 48h following the decline of the bloom. Synchrotron x-ray fluorescence (SXRF) analyses of individual cells showed that constituent elements were lost from sinking cells at notably different rates (Fig. 3). Phosphorus was rapidly released from sinking cells, with mean P quotas decreasing 55% and 73% from surface values by 100m and 200m, respectively (30). However, only 25% of cellular Fe was lost from cells sinking through the upper 200m, while 61% of cellular Ni was remineralized. This supports the story told by the bulk biogeochemical data from the North Atlantic: Ni is remineralized largely to a similar degree as P, while Fe is lost more slowly from sinking biogenic material.

Application of microanalytical techniques such as SXRF can be combined with bulk approaches to further advance understanding of subsurface metal remineralization and cycling. In FeCycle II, Fe:P of sinking A. glacialis cells increased, on average, only 2.3-fold in the upper 200m, while Fe:P in bulk particulate matter increased more than 13-fold (30). This indicates that the behavior of sinking cells was not representative of the full particle assemblage. Iron and P were likely more completely decoupled in sinking fecal pellets and detrital material (which appears to have contributed significantly to the particulate Fe pool during FeCycle II; 31) than in intact sinking cells. Further application of this approach will allow us to not only distinguish between the behavior of biogenic and lithogenic fractions (Fig. 1), but potentially also between detrital particles. By considering metals such as Mn that are prone to oxidation and scavenging in the subsurface ocean (32, 33), it may also be possible to separate abiotic scavenging from net biological remineralization (Fig. 1). Additionally, 2D (and potentially 3D) mapping of elements within cells and particles also provides information about the spatial and potentially chemical associations of elements with particles (30, 34).

The GEOTRACES program is generating unprecedented data, both in terms of quality and quantity, regarding the cycling of bioactive trace metals in the ocean. Syntheses of these data, and integration of insights from novel microanalytical tools, as well as transcriptomic and proteomic approaches, are resulting in substantial advances in our understanding of metal biogeochemistry. No longer are we limited to a few painstakingly collected dissolved metal profiles. There is now painstakingly collected full-depth coverage of most ocean basins, including in many cases dissolved and particulate fractions of nearly all biogenic elements, enabling testing of early hypotheses about trace metal cycling and parameterization of these processes into next-generation ocean biogeochemical models.

Authors

Benjamin S. Twining, Daniel C. Ohnemus, Renee L. Torrie (Bigelow Laboratory for Ocean Sciences)

Acknowledgments

This work was funded by NSF grant OCE-1232814 to BST. RLT was funded by NSF REU grant 1460861 to Bigelow Laboratory for Ocean Sciences.

References

1. A. C. Redfield, in James Johnstone Memorial Volume, R. J. Daniel, Ed. (Liverpool University Press, 1934), pp. 176-192.
2. A. C. Redfield, Amer. Scientist 46, 205-221 (1958).
3. E. A. Boyle, J. M. Edmond, Nature 253, 107-109 (1975).
4. E. A. Boyle, F. Sclater, J. M. Edmond, Nature 263, 42-44 (1976).
5. K. W. Bruland, Earth Plan. Sci. Lett. 47, 176-198 (1980).
6. J. R. R. Frausto da Silva, R. J. P. Williams, The Biological Chemistry of the Elements: the inorganic chemistry of life. (Oxford University Press, Oxford, ed. 2nd, 2001), pp. 575.
7. F. M. M. Morel, R. J. M. Hudson, in Chemical Processes in Lakes, W. Stumm, Ed. (John Wiley & Sons, New York, 1985), pp. 251-281.
8. K. W. Bruland, J. R. Donat, D. A. Hutchins, Limnol. Oceanogr. 36, 1555-1577 (1991).
9. T. Y. Ho et al., J. Phycol. 39, 1145-1159 (2003).
10. W. G. Sunda, Marine Chem. 57, 169-172 (1997).
11. W. G. Sunda, S. A. Huntsman, Limnol. Oceanogr. 40, 132-137 (1995).
12. W. G. Sunda, S. A. Huntsman, Limnol. Oceanogr. 40, 1404-1417 (1995).
13. C. M. Moore et al., Nature Geosci. 6, 701-710 (2013).
14. R. Collier, J. Edmond, Prog. Oceanogr. 13, 113-199 (1984).
15. F. M. M. Morel, Geobiol. 6, 318-324 (2008).
16. M. Whitfield, D. R. Turner, in Aquatic Surface Chemistry: Chemical Processes at the Particle-Water Interface, W. Stumm, Ed. (John Wiley & Sons, Inc., 1987), pp. 457-493.
17. K. W. Bruland, M. C. Lohan, in The Oceans and Marine Geochemistry: Treatise on Geochemistry, H. Elderfield, Ed. (Elsevier, Oxford, 2003), pp. 23-47.
18. P. W. Boyd, M. J. Ellwood, Nature Geosci. 3, 675-682 (2010).
19. D. C. Ohnemus, P. J. Lam, Cycling of lithogenic marine particles in the US GEOTRACES North Atlantic Transect. Deep-Sea Res. II 116, 282-302 (2015).
20. P. W. Boyd et al., Limnol. Oceanogr. 55, 1271-1288 (2010).
21. R. D. Frew et al., Glob. Biogeochem. Cycles 20, GB1S93, doi:10.1029/2005GB002558 (2006).
22. C. H. Lamborg, K. O. Buesseler, P. J. Lam, Deep-Sea Res. II 55, 1564-1577 (2008).
23. W. J. Jenkins et al., Deep-Sea Res. II 116, 6-20 (2015).
24. M. Gledhill, K. N. Buck, Frontiers Microbiol. 3, doi: 10.3389/fmicb.2012.00069 (2012).
25. J. W. Moffett, J. Ho, Geochim. Cosmochim. Acta 60, 3415-3424 (1996).
26. A. E. Noble et al., Limnol. Oceanogr. 57, 989-1010 (2012).
27. B. S. Twining et al., Glob. Biogeochem. Cycles 26, GB4001, doi:4010.1029/2011GB004233 (2012).
28. P. J. Lam et al., Prog. Oceanogr. 133, 32-42 (2015).
29. P. W. Boyd et al., Geophys. Res. Lett. 39, doi:10.1029/2012GL053448 (2012).
30. B. S. Twining et al., Limnol. Oceanogr. 59, 689-704 (2014).
31. A. L. King et al., Biogeosci. 9, 667-687 (2012).
32. J. P. Cowen, K. W. Bruland, Deep-Sea Res. 32, 253-272 (1985).
33. D. C. Ohnemus et al., Limnol. Oceanogr. In press (2016).
34. J. Nuester, S. Vogt, B. S. Twining, J. Phycol. 48, 626-634 (2012). 35. B. S. Twining et al., Prog. Oceanogr. 137, 261-283 (2015).
36. M. Hatta et al., Deep-Sea Res. II 2015, 117-129 (2015).
37. S. Roshan, J. Wu, Glob. Biogeochem. Cycles 29, 2082-2094 (2015).
38. E. Mawji et al. Marine Chem. 177, 1-8 (2015).

Filter by Keyword

abundance acidification additionality advection africa air-sea air-sea interactions algae alkalinity allometry ammonium AMO AMOC anoxic Antarctic Antarctica anthro impacts anthropogenic carbon anthropogenic impacts appendicularia aquaculture aquatic continuum aragonite saturation arctic Argo argon arsenic artificial seawater Atlantic atmospheric CO2 atmospheric nitrogen deposition authigenic carbonates autonomous platforms bacteria bathypelagic BATS BCG Argo benthic bgc argo bio-go-ship bio-optical bioavailability biogeochemical cycles biogeochemical models biogeochemistry Biological Essential Ocean Variables biological pump biophysics bloom blue carbon bottom water boundary layer buffer capacity C14 CaCO3 calcification calcite carbon carbon-climate feedback carbon-sulfur coupling carbonate carbonate system carbon budget carbon cycle carbon dioxide carbon export carbon fluxes carbon sequestration carbon storage Caribbean CCA CCS changing marine chemistry changing marine ecosystems changing marine environments changing ocean chemistry chemical oceanographic data chemical speciation chemoautotroph chesapeake bay chl a chlorophyll circulation CO2 coastal and estuarine coastal darkening coastal ocean cobalt Coccolithophores commercial community composition competition conservation cooling effect copepod copepods coral reefs CTD currents cyclone daily cycles data data access data assimilation database data management data product Data standards DCM dead zone decadal trends decomposers decomposition deep convection deep ocean deep sea coral denitrification deoxygenation depth diatoms DIC diel migration diffusion dimethylsulfide dinoflagellate dinoflagellates discrete measurements distribution DOC DOM domoic acid DOP dust DVM ecology economics ecosystem management ecosystems eddy Education EEZ Ekman transport emissions ENSO enzyme equatorial current equatorial regions ESM estuarine and coastal carbon fluxes estuary euphotic zone eutrophication evolution export export fluxes export production extreme events faecal pellets fecal pellets filter feeders filtration rates fire fish Fish carbon fisheries fishing floats fluid dynamics fluorescence food webs forage fish forams freshening freshwater frontal zone functional role future oceans gelatinous zooplankton geochemistry geoengineering geologic time GEOTRACES glaciers gliders global carbon budget global ocean global warming go-ship grazing greenhouse gas greenhouse gases Greenland ground truthing groundwater Gulf of Maine Gulf of Mexico Gulf Stream gyre harmful algal bloom high latitude human food human impact human well-being hurricane hydrogen hydrothermal hypoxia ice age ice cores ice cover industrial onset inland waters in situ inverse circulation ions iron iron fertilization iron limitation isotopes jellies katabatic winds kelvin waves krill kuroshio lab vs field land-ocean continuum larvaceans lateral transport LGM lidar ligands light light attenuation lipids low nutrient machine learning mangroves marine carbon cycle marine heatwave marine particles marine snowfall marshes mCDR mechanisms Mediterranean meltwater mesopelagic mesoscale mesoscale processes metagenome metals methane methods microbes microlayer microorganisms microplankton microscale microzooplankton midwater mitigation mixed layer mixed layers mixing mixotrophs mixotrophy model modeling model validation mode water molecular diffusion MPT MRV multi-decade n2o NAAMES NCP nearshore net community production net primary productivity new ocean state new technology Niskin bottle nitrate nitrogen nitrogen cycle nitrogen fixation nitrous oxide north atlantic north pacific North Sea nuclear war nutricline nutrient budget nutrient cycles nutrient cycling nutrient limitation nutrients OA observations ocean-atmosphere ocean acidification ocean acidification data ocean alkalinity enhancement ocean carbon storage and uptake ocean carbon uptake and storage ocean color ocean modeling ocean observatories ocean warming ODZ oligotrophic omics OMZ open ocean optics organic particles oscillation outwelling overturning circulation oxygen pacific paleoceanography PAR parameter optimization parasite particle flux particles partnerships pCO2 PDO peat pelagic PETM pH phenology phosphate phosphorus photosynthesis physical processes physiology phytoplankton PIC piezophilic piezotolerant plankton POC polar polar regions policy pollutants precipitation predation predator-prey prediction pressure primary productivity Prochlorococcus productivity prokaryotes proteins pteropods pycnocline radioisotopes remineralization remote sensing repeat hydrography residence time resource management respiration resuspension rivers rocky shore Rossby waves Ross Sea ROV salinity salt marsh satellite scale seafloor seagrass sea ice sea level rise seasonal seasonality seasonal patterns seasonal trends sea spray seawater collection seaweed secchi sediments sensors sequestration shelf ocean shelf system shells ship-based observations 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

Copyright © 2025 - OCB Project Office, Woods Hole Oceanographic Institution, 266 Woods Hole Rd, MS #25, Woods Hole, MA 02543 USA Phone: 508-289-2838  •  Fax: 508-457-2193  •  Email: ocb_news@us-ocb.org

link to nsflink to noaalink to WHOI

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.