Deborah K. Steinberg

Eddy/Wind Interactions Stimulate Extraordinary Mid-Ocean Plankton Blooms

Episodic eddy-driven upwelling may supply a significant fraction of the nutrients required to sustain primary productivity of the subtropical ocean. New observations in the northwest Atlantic reveal that, although plankton blooms occur in both cyclones and mode-water eddies, the biological responses differ. Mode-water eddies can generate extraordinary diatom biomass and primary production at depth, relative to the time series near Bermuda. These blooms are sustained by eddy/wind interactions, which amplify the eddy-induced upwelling. In contrast, eddy/wind interactions dampen eddy-induced upwelling in cyclones. Carbon export inferred from oxygen anomalies in eddy cores is one to three times as much as annual new production for the region.

Overview of the US JGOFS Bermuda Atlantic Time-series Study (BATS): a decade-scale look at ocean biology and biogeochemistry

The Bermuda Atlantic Time-series Study (BATS) commenced monthly sampling in October 1988 as part of the US Joint Global Ocean Flux Study (JGOFS) program. The goals of the US JGOFS time-series research are to better understand the basic processes that control ocean biogeochemistry on seasonal to decadal time-scales, determine the role of the oceans in the global carbon budget, and ultimately improve our ability to predict the effects of climate change on ecosystems. The BATS program samples the ocean on a biweekly to monthly basis, a strategy that resolves major seasonal patterns and interannual variability. The core cruises last 4-5 d during which hydrography, nutrients, particle flux, pigments and primary production, bacterioplankton abundance and production, and often complementary ancillary measurements are made. This overview focuses on patterns in ocean biology and biogeochemistry over a decade at the BATS site, concentrating on seasonal and interannual changes in community structure, and the physical forcing and other factors controlling the temporal dynamics. Significant seasonal and interannual variability in phytoplankton and bacterioplankton production, biomass, and community structure exists at BATS. No strong relationship exists between primary production and particle flux during the 10 yr record, with the relationship slightly improved by applying an artificial lag of 1 week between production and flux. The prokaryotic picoplankton regularly dominate the phytoplankton community; diatom blooms are rare but occur periodically in the BATS time series. The increase in Chi a concentrations during bloom periods is due to increases by most of the taxa present, rather than by any single group, and there is seasonal succession of phytoplankton. The bacterioplankton often dominate the living biomass, indicating the potential to consume large amounts of carbon and play a major ecological role within the microbial food web. Bacterial biomass, production, and specific growth rates are highest during summer. Size structure and composition of the plankton community may be an important factor controlling the quality of dissolved organic matter produced and could affect production of bacterioplankton biomass. Larger heterotrophic plankton play an integral role in the flux of material out of the euphotic zone at BATS. Protozoans are abundant and can constitute a sizable component of sinking flux. Zooplankton contribute significantly to flux via production of rapidly sinking fecal pellets, and vertically migrating zooplankton can actively transport a significant amount of dissolved organic and inorganic carbon and nitrogen to deep water. An important question that remains to be further addressed at BATS is how larger climatological events drive some of the interannual variability in the biogeochemistry

An assessment of the use of sediment traps for estimating upper ocean particle fluxes

This review provides an assessment of sediment trap accuracy issues by gathering data to address trap hydrodynamics, the problem of zooplankton “swimmers,” and the solubilization of material after collection. For each topic, the problem is identified, its magnitude and causes reviewed using selected examples, and an update on methods to correct for the potential bias or minimize the problem using new technologies is presented. To minimize hydrodynamic biases due to flow over the trap mouth, the use of neutrally buoyant sediment traps is encouraged. The influence of swimmers is best minimized using traps that limit zooplankton access to the sample collection chamber. New data on the impact of different swimmer removal protocols at the US time-series sites HOT and BATS are compared and shown to be important. Recent data on solubilization are compiled and assessed suggesting selective losses from sinking particles to the trap supernatant after collection, which may alter both fluxes and ratios of elements in long term and typically deeper trap deployments. Different methods are needed to assess shallow and short- term trap solubilization effects, but thus far new incubation experiments suggest these impacts to be small for most elements. A discussion of trap calibration methods reviews independent assessments of flux, including elemental budgets, particle abundance and flux modeling, and emphasizes the utility of U-Th radionuclide calibration methods.

Zooplankton vertical migration and the active transport of dissolved organic and inorganic carbon in the Sargasso Sea

The least known component of the “biological pump” is the active transport of carbon and nutrients by diel vertical migration of zooplankton. We measured CO2 respiration and dissolved organic carbon (DOC) excretion by individual species of common vertically migrating zooplankton at the US JGOFS Bermuda Atlantic Time-series Study (BATS) station. The inclusion of DOC excretion in this study builds on published research on active transport by respiration of inorganic carbon and allows a direct assessment of the role of zooplankton in the production of dissolved organic matter used in midwater microbial processes. On average, excretion of DOC makes up 24% (range=5–42%) of the total C metabolized (excreted+respired) and could represent a significant augmentation to the vertical flux that has already been documented for respiratory CO2 flux by migrant zooplankton. Migratory fluxes were compared to other transport processes at BATS. Estimates of combined active transport of CO2 and DOC by migrators at BATS averaged 7.8% and reached 38.6% of mean sinking POC flux at 150 m, and reached 71.4% of mean sinking POC flux at 300 m. DOC export by migrator excretion averaged 1.9% and reached 13.3% of annual DOC export by physical mixing at this site. During most of the year when deep mixing does not occur, diel migration by zooplankton could provide a supply of DOC to the deeper layers that is available for use by the microbial community. A carbon budget comparing migrant zooplankton transport to the balance of fluxes in the 300–600 m depth strata at BATS shows on average that the total migrant flux supplies 37% of the organic carbon remineralized in this layer, and that migrant DOC flux is more than 3 times the DOC flux gradient by diapycnal mixing. New estimates of active transport of both organic and inorganic carbon by migrants may help resolve observed imbalances in the C budget at BATS, but the magnitude is highly dependent on the biomass of the migrating community.

Jellyfish blooms result in a major microbial respiratory sink of carbon in marine systems.

Jellyfish blooms occur in many estuarine and coastal regions and may be increasing in their magnitude and extent worldwide. Voracious jellyfish predation impacts food webs by converting large quantities of carbon (C), fixed by primary producers and consumed by secondary producers, into gelatinous biomass, which restricts C transfer to higher trophic levels because jellyfish are not readily consumed by other predators. In addition, jellyfish release colloidal and dissolved organic matter (jelly-DOM), and could further influence the functioning of coastal systems by altering microbial nutrient and DOM pathways, yet the links between jellyfish and bacterioplankton metabolism and community structure are unknown. Here we report that jellyfish released substantial quantities of extremely labile C-rich DOM, relative to nitrogen (25.6 ± 31.6 C:1N), which was quickly metabolized by bacterioplankton at uptake rates two to six times that of bulk DOM pools. When jelly-DOM was consumed it was shunted toward bacterial respiration rather than production, significantly reducing bacterial growth efficiencies by 10% to 15%. Jelly-DOM also favored the rapid growth and dominance of specific bacterial phylogenetic groups (primarily γ-proteobacteria) that were rare in ambient waters, implying that jelly-DOM was channeled through a small component of the in situ microbial assemblage and thus induced large changes in community composition. Our findings suggest major shifts in microbial structure and function associated with jellyfish blooms, and a large detour of C toward bacterial CO2 production and away from higher trophic levels. These results further suggest fundamental transformations in the biogeochemical functioning and biological structure of food webs associated with jellyfish blooms.

Winter and spring controls on the summer food web of the coastal West Antarctic Peninsula

Understanding the mechanisms by which climate variability affects multiple trophic levels in food webs is essential for determining ecosystem responses to climate change. Here we use over two decades of data collected by the Palmer Long Term Ecological Research program (PAL-LTER) to determine how large-scale climate and local physical forcing affect phytoplankton, zooplankton and an apex predator along the West Antarctic Peninsula (WAP). We show that positive anomalies in chlorophyll-a (chl-a) at Palmer Station, occurring every 4-6 years, are constrained by physical processes in the preceding winter/spring and a negative phase of the Southern Annular Mode (SAM). Favorable conditions for phytoplankton included increased winter ice extent and duration, reduced spring/summer winds, and increased water column stability via enhanced salinity-driven density gradients. Years of positive chl-a anomalies are associated with the initiation of a robust krill cohort the following summer, which is evident in Adélie penguin diets, thus demonstrating tight trophic coupling. Projected climate change in this region may have a significant, negative impact on phytoplankton biomass, krill recruitment and upper trophic level predators in this coastal Antarctic ecosystem.

Prediction of the Export and Fate of Global Ocean Net Primary Production: The EXPORTS Science Plan

Ocean ecosystems play a critical role in the Earth’s carbon cycle and the quantification of their impacts for both present conditions and for predictions into the future remains one of the greatest challenges in oceanography. The goal of the EXport Processes in the Ocean from Remote Sensing (EXPORTS) Science Plan is to develop a predictive understanding of the export and fate of global ocean net primary production (NPP) and its implications for present and future climates. The achievement of this goal requires a quantification of the mechanisms that control the export of carbon from the euphotic zone as well as its fate in the underlying “twilight zone” where some fraction of exported carbon will be sequestered in the ocean’s interior on time scales of months to millennia. Here we present a measurement / synthesis / modeling framework aimed at quantifying the fates of upper ocean NPP and its impacts on the global carbon cycle based upon the EXPORTS Science Plan. The proposed approach will diagnose relationships among the ecological, biogeochemical and physical oceanographic processes that control carbon cycling across a range of ecosystem and carbon cycling states leading to advances in satellite diagnostic and numerical prognostic models. To collect these data, a combination of ship and robotic field sampling, satellite remote sensing and numerical modeling is proposed which enables the sampling of the many pathways of NPP export and fates. This coordinated, process-oriented approach has the potential to foster new insights on ocean carbon cycling that maximizes its societal relevance through the achievement of research goals of many international research agencies and will be a key step towards our understanding of the Earth as an integrated system.

Increased Feeding and Nutrient Excretion of Adult Antarctic Krill, Euphausia superba, Exposed to Enhanced Carbon Dioxide (CO2)

Ocean acidification has a wide-ranging potential for impacting the physiology and metabolism of zooplankton. Sufficiently elevated CO2 concentrations can alter internal acid-base balance, compromising homeostatic regulation and disrupting internal systems ranging from oxygen transport to ion balance. We assessed feeding and nutrient excretion rates in natural populations of the keystone species Euphausia superba (Antarctic krill) by conducting a CO2 perturbation experiment at ambient and elevated atmospheric CO2 levels in January 2011 along the West Antarctic Peninsula (WAP). Under elevated CO2 conditions (∼672 ppm), ingestion rates of krill averaged 78 µg C individual−1 d−1 and were 3.5 times higher than krill ingestion rates at ambient, present day CO2 concentrations. Additionally, rates of ammonium, phosphate, and dissolved organic carbon (DOC) excretion by krill were 1.5, 1.5, and 3.0 times higher, respectively, in the high CO2 treatment than at ambient CO2 concentrations. Excretion of urea, however, was ∼17% lower in the high CO2 treatment, suggesting differences in catabolic processes of krill between treatments. Activities of key metabolic enzymes, malate dehydrogenase (MDH) and lactate dehydrogenase (LDH), were consistently higher in the high CO2 treatment. The observed shifts in metabolism are consistent with increased physiological costs associated with regulating internal acid-base equilibria. This represents an additional stress that may hamper growth and reproduction, which would negatively impact an already declining krill population along the WAP.

Zooplankton and the Ocean Carbon Cycle

Marine zooplankton comprise a phylogenetically and functionally diverse assemblage of protistan and metazoan consumers that occupy multiple trophic levels in pelagic food webs. Within this complex network, carbon flows via alternative zooplankton pathways drive temporal and spatial variability in production-grazing coupling, nutrient cycling, export, and transfer efficiency to higher trophic levels. We explore current knowledge of the processing of zooplankton food ingestion by absorption, egestion, respiration, excretion, and growth (production) processes. On a global scale, carbon fluxes are reasonably constrained by the grazing impact of microzooplankton and the respiratory requirements of mesozooplankton but are sensitive to uncertainties in trophic structure. The relative importance, combined magnitude, and efficiency of export mechanisms (mucous feeding webs, fecal pellets, molts, carcasses, and vertical migrations) likewise reflect regional variability in community structure. Climate change is expected to broadly alter carbon cycling by zooplankton and to have direct impacts on key species.

Nitrogen Consumption and Metabolism in Marine Zooplankton

Publisher Summary This chapter focuses on the important role that zooplankton play in nitrogen cycling by following the path of the food from ingestion to all aspects and stages of metabolism. It begins with a discussion of N consumption, including the nitrogenous composition of available food types, rates of N consumption, and effects of food quality. The various aspects of N metabolism, emphasizing the importance to biogeochemical cycling and coupling with other major elements (carbon, phosphorus) are also discussed. Considerable progress in our understanding of the role of marine zooplankton in nitrogen cycling has been made in the last two decades. Studies are now at the stage where large data sets are being compiled, and it is possible, for example, to reasonably predict ammonia excretion as a function of body mass for a given organism and temperature using regression equations—and avoid doing experimental incubations. These individual, size-specific excretion rates can be scaled up to a community rate if total biomass and size structure of the community is known. A common theme throughout this review is that more information is needed on the role of marine protozoa, gelatinous zooplankton, and of taxa living below the euphotic zone in N consumption and metabolism. Consumption of phytoplankton and bacteria by microzooplankton are key processes in N transformations in the ocean.