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Climate-driven pelagification of marine food webs: Implications for marine fish populations

Posted by mmaheigan

· Friday, January 22nd, 2021

Global warming changes the conditions for all ocean life, with wide-ranging consequences. It is particularly difficult to predict the impact of climate change on fish because fish production is conditioned on both temperature and food resource (zooplankton and benthic organisms) changes. Climate change projections from Earth system models show a negative amplification of changes in global ocean net primary production (NPP), with an approximate doubling of production decreases from net primary producers to mesozooplankton. This “trophic amplification” continues up the marine food web to fishes. A new study published in Frontiers in Marine Science illustrates this amplification clearly when fishes are defined by their maximum body size, which describes their position in the food web (Figure 1a). However, decreases in globally integrated biomass and production were not limited to differences in size alone. Importantly, reduced abundances also varied by fish functional type (Figure 1b).

Figure 1: a) Percent change in net primary production (NPP), mesozooplankton (MesoZ) production, all medium (M) fishes, and all large (L) fishes from Historic (1951-2000) to the RCP 8.5 Projection (2051-2100). b) Percent change in production of forage fish, large pelagic fish, demersal fish, and benthic invertebrates in Projection (2051-2100) from Historic (1951-2000). c) Absolute change in the ratio of zooplankton production to seafloor detrital flux as the difference of the Projection (2051-2100) from the Historic (1951-2000). d) Percent change in zooplankton production (dashed grey), percent change in seafloor detrital flux (solid grey), and absolute change in the ratio of their means during the Historic and Projection time periods relative to 1951.

Despite the “pelagification” of marine food webs caused by unequal decreases in secondary production (Figure 1d) and subsequent increases in pelagic zooplankton production relative to seafloor detritus production (Figure 1c,d), large pelagic fish (e.g., tunas and billfishes) suffered the greatest declines and the highest degree of projection uncertainty. The result was a shift from benthic-based ecosystems historically dominated by large demersal fish (e.g., cods and flounders) towards pelagic-based ones dominated by smaller forage fish (e.g., sardines and herring). Any positive impacts of the pelagification of food resources on large pelagic fish were overwhelmed by the negative impacts of the overall reduction in global productivity, compounded by warming-induced increases in metabolic demands. Both the degree of change in the productivity of large pelagic fish and the magnitude of trophic amplification were sensitive to the temperature dependence of metabolic rates. Thus, better constraints are needed on empirical estimates of the effect of temperature on physiological rates to project the impacts of climate change on fish biomass and marine ecosystem structure.

Ocean fish harvests currently supply ~15% of global protein demand. Reduced primary production will decrease the total amount of fish available to harvest for human food, while the pelagification of ecosystems could require large and expensive structural modifications to fisheries, including gear, location, regional and international management plans, consumer demands, and market values.

Figure caption: a) Percent change in net primary production (NPP), mesozooplankton (MesoZ) production, all medium (M) fishes, and all large (L) fishes from Historic (1951-2000) to the RCP 8.5 Projection (2051-2100). b) Percent change in production of forage fish, large pelagic fish, demersal fish, and benthic invertebrates in Projection (2051-2100) from Historic (1951-2000). c) Absolute change in the ratio of zooplankton production to seafloor detrital flux as the difference of the Projection (2051-2100) from the Historic (1951-2000). d) Percent change in zooplankton production (dashed grey), percent change in seafloor detrital flux (solid grey), and absolute change in the ratio of their means during the Historic and Projection time periods relative to 1951.

Authors:

Colleen M. Petrik, Texas A&M University

Charles A. Stock, Geophysical Fluid Dynamics Laboratory

Ken H. Andersen, Technical University of Denmark

P. Daniël van Denderen, International Council for the Exploration of the Seas

James R. Watson, Oregon State University

Austral summer vertical migration patterns in Antarctic zooplankton

Posted by mmaheigan

· Thursday, October 15th, 2020

Sunrise and sunset are the main cues driving zooplankton diel vertical migration (DVM) throughout the world’s oceans. These marine animals balance the trade-off between feeding in surface waters at night and avoiding predation during the day at depth. Near-constant daylight during polar summer was assumed to dampen these daily migrations. In a recent paper published in Deep-Sea Research I, authors assessed austral summer DVM patterns for 15 taxa over a 9-year period. Despite up to 22 hours of sunlight, a diverse array of zooplankton – including copepods, krill, pteropods, and salps – continued DVM.

Figure caption: Mean day (orange) and night (blue) abundance of (A) the salp Salpa thompsoni, (B) the krill species Thysanoessa macrura, (C) the pteropod Limacina helicina, and (D) chaetognaths sampled at discrete depth intervals from 0-500m. Horizontal dashed lines indicate weighted mean depth (WMD). N:D is the night to day abundance ratio for 0-150 m. Error bars indicate one standard error. Sample size n = 12 to 22. Photos by Larry Madin, Miram Gleiber, and Kharis Schrage.

The Palmer Antarctica Long-Term Ecological Research (LTER) Program conducted this study using a MOCNESS (Multiple Opening/Closing Net and Environmental Sensing System) to collect depth-stratified samples west of the Antarctic Peninsula. The depth range of migrations during austral summer varied across taxa and with daylength and phytoplankton biomass and distribution. While most taxa continued some form of DVM, others (e.g., carnivores and detritivores) remained most abundant in the mesopelagic zone, regardless of photoperiod, which likely impacted the attenuation of vertical carbon flux. Given the observed differences in vertical distribution and migration behavior across taxa, ongoing changes in Antarctic zooplankton assemblages will likely impact carbon export pathways. More regional, taxon-specific studies such as this are needed to inform efforts to model zooplankton contributions to the biological carbon pump.

Authors:
John Conroy (VIMS, William & Mary)
Deborah Steinberg (VIMS, William & Mary)
Patricia Thibodeau (VIMS, William & Mary; currently University of Rhode Island)
Oscar Schofield (Rutgers University)

Will global change “stress out” ocean DOC cycling?

Posted by mmaheigan

· Tuesday, September 29th, 2020

The dissolved organic carbon (DOC) pool is vital for the functioning of marine ecosystems. DOC fuels marine food webs and is a cornerstone of the earth’s carbon cycle. As one of the largest pools of organic matter on the planet, disruptions to marine DOC cycling driven by climate and environmental global changes can impact air-sea CO2 exchange, with the added potential for feedbacks on Earth’s climate system.

Figure 1. Simplified view of major dissolved organic carbon (DOC) sources (black text) and sinks (yellow text) in the ocean.

Since DOC cycling involves multiple processes acting concurrently over a range of time and space scales, it is especially challenging to characterize and quantify the influence of global change. In a recent review paper published in Frontiers in Marine Science, the authors synthesize impacts of global change-related stressors on DOC cycling such as ocean warming, stratification, acidification, deoxygenation, glacial and sea ice melting, inflow from rivers, ocean circulation and upwelling, and atmospheric deposition. While ocean warming and acidification are projected to stimulate DOC production and degradation, in most regions, the outcomes for other key climate stressors are less clear, with much more regional variation. This synthesis helps advance our understanding of how global change will affect the DOC pool in the future ocean, but also highlights important research gaps that need to be explored. These gaps include for example a need for studies that allow to understand the adaptation of degradation/production pathways to global change stressors, and their cumulative impacts (e.g. temperature with acidification).

Authors:
C. Lønborg (Aarhus University)
C. Carreira (CESAM, Universidade de Aveiro)
Tim Jickells (University of East Anglia)
X.A. Álvarez-Salgado (CSIC, Instituto de Investigacións Mariñas)

Turning a spotlight on grazing

Posted by mmaheigan

· Thursday, July 23rd, 2020

Microscopic plankton in the surface ocean make planet Earth habitable by generating oxygen and forming the basis of marine food webs, yielding harvestable protein. For over 100 years, oceanographers have tried to ascertain the physical, chemical, and biological processes governing phytoplankton blooms. Zooplankton grazing of phytoplankton is the single largest loss process for primary production, but empirical grazing data are sparse and thus poorly constrained in modeling frameworks, including assessments of global elemental cycles, cross-ecosystem comparisons, and predictive efforts anticipating future ocean ecosystem function. As sunlight decays exponentially with depth, upper-ocean mixing creates dynamic light environments with predictable effects on phytoplankton growth but unknown consequences for grazing.

Figure caption: Rates (d⁻¹) of phytoplankton growth (μ), grazing mortality (g), and biomass accumulation (r) under four mixed layer scenarios simulated using light as a proxy of (a) sustained deep mixing, (b) rapid shoaling, (c) sustained shallow mixing, and (d) rapid mixed layer deepening. Error bars represent one standard deviation of the mean of duplicate experiments. Grazing was measured but not detected in the sustained deep mixing and rapid shoaling conditions, denoted with x.

Using data from a spring cruise in the North Atlantic, authors of a recent study published in Limnology & Oceanography compared the influences of microzooplankton predation and fluctuations in light availability—representative of a mixing water column—on phytoplankton standing stock. Data from at-sea incubations and light manipulation experiments provide evidence that phytoplankton’s instantaneous and zooplankton’s delayed responses to light fluctuations are key modulators of the balance between phytoplankton growth and grazing rates (Figure 1). These results suggest that light is a potential, remotely retrievable predictor of when and where in the ocean zooplankton grazing may represent an important loss term of phytoplankton production. If broadly verified, this approach could be used to systematically assess sparsely measured grazing across spatial and temporal gradients in representative regions of the ocean. Such data will be essential for enhancing our predictive capacity of ocean food web function, global biogeochemical cycles and the many derived processes, including fisheries production and the flow of carbon through the oceans.

Authors:
Françoise Morison (University of Rhode Island)
Gayantonia Franzè (University of Rhode Island, currently Institute of Marine Research, Norway)
Elizabeth Harvey (University of Georgia, currently University of New Hampshire)
Susanne Menden-Deuer (University of Rhode Island)

Modern OMZ copepod dynamics provide analog for future oceans

Posted by mmaheigan

· Thursday, July 23rd, 2020

Global warming increases ocean deoxygenation and expands the oxygen minimum zone (OMZ), which has implications for major zooplankton groups like copepods. Reduced oxygen levels may impact individual copepod species abundance, vertical distribution, and life history strategy, which is likely to perturb intricate oceanic food webs and export processes. In a study recently published in Biogeosciences, authors conducted vertically-stratified day and night MOCNESS tows (0-1000 m) during four cruises (2007-2017) in the Eastern Tropical North Pacific, sampling hydrography and copepod distributions in four locations with different water column oxygen profiles and OMZ intensity (i.e. lowest oxygen concentration and its vertical extent in a profile). Each copepod species exhibited a different vertical distribution strategy and physiology associated with oxygen profile variability. The study identified sets of species that (1) changed their vertical distributions and maximum abundance depth associated with the depth and intensity of the OMZ and its oxycline inflection points, (2) shifted their diapause depth, (3) adjusted their diel vertical migration, especially the nighttime upper depth, or (4) expanded or contracted their depth range within the mixed layer and upper part of the thermocline in association with the thickness of the aerobic epipelagic zone (habitat compression concept) (Figure 1). Distribution depths for some species shifted by 10’s to 100’s of meters in different situations, which also had metabolic (and carbon flow) implications because temperature decreased with depth. This observed present-day variability may provide an important window into how future marine ecosystems will respond to deoxygenation.

Figure caption: Schematic diagram showing how future OMZ expansion may affect zooplankton distributions, based on present-day responses to OMZ variability. The dashed line indicates diel vertical migration (DVM) and highlights the shoaling of the nighttime depth as the aerobic habitat is compressed. The lower oxycline community and the diapause layer for some species, associated with a specific oxygen concentration, may deepen as the OMZ expands.

Authors:
Karen F. Wishner (University of Rhode Island)
Brad Seibel (University of South Florida)
Dawn Outram (University of Rhode Island)

Autonomous platforms yield new insights on North Atlantic bloom phenology

Posted by mmaheigan

· Wednesday, April 22nd, 2020

Phytoplankton produces organic carbon, which serves as a major energy source in marine food webs and plays an important role in the global carbon cycle. Studies of phytoplankton seasonal timing (phenology) have been a major focus in oceanography, especially in the subpolar North Atlantic region, where massive increases in phytoplankton biomass (blooms) occur during the winter-spring transition.

Figure 1. Panel a: Each line represents the trajectory of a profiling Argo float deployed during the North Atlantic Aerosols and Marine Ecosystems Study (NAAMES) expeditions (12 total); the initial float deployment location is denoted by a filled circle. The bar chart (inset right bottom) indicates float deployment durations. Panel b: Seasonal climatologies of C_phyto (green), µ (blue), l (red), and r (grey) from Argo floats for all 4 regions (D1-D4 as indicated on map in Panel a).

Many hypotheses based on data from shipboard discrete sampling or satellite remote sensing have been proposed to explain drivers of phytoplankton bloom formation and dynamics. However, discrete shipboard sampling limits both spatial and temporal coverage, and satellite approaches cannot provide direct information at depth. To address this gap in spatiotemporal coverage, a recent study in Frontiers in Marine Science, applied bio-optical measurements from 12 Argo profiling floats to study the year-round phytoplankton phenology in a north-south section of the western North Atlantic Ocean (40° N to 60° N). The authors calculated phytoplankton division rate (µ), loss rate (l), and carbon accumulation rate (r) using the Argo-based Chlorophyll-a (Chl) and phytoplankton carbon (C_phyto) estimates. Latitudinally varying phytoplankton dynamics were observed, with a higher (and later) C_phyto peak in the north, and stronger μ–r decoupling and increased proportion of winter to total annual production in the south (Figure 1). Seasonal phenology patterns arise from interactions between “bottom-up” (e.g., resources for growth) and “top-down” (e.g., grazing, mortality) factors that involve both biological and physical drivers. The Argo float data are consistent with the disturbance recovery hypothesis (DRH) over a full annual cycle. Float-based mixed layer phytoplankton phenology observations were comparable to satellite remote sensing observations. In a data-model comparison, outputs from an eddy-resolving ocean simulation only reproduced some of the observed phytoplankton phenology, indicating possible biases in the simulated physical forcing, turbulent dynamics, and biophysical interactions.

In addition to seasonal patterns in the mixed layer, float-based measurements provide information on the vertical distribution of physical and biogeochemical quantities and therefore are complementary to the satellite measurements. This powerful combination of observing assets enhances spatiotemporal coverage, thus enabling us to better observe, compare, model, and predict seasonal phytoplankton dynamics in the subpolar North Atlantic.

Authors:
Bo Yang (University of Virginia)
Emmanuel S. Boss (University of Maine)
Nils Haëntjens (University of Maine)
Matthew C. Long (National Center for Atmospheric Research)
Michael J. Behrenfeld (Oregon State University)
Rachel Eveleth (Oberlin College)
Scott C. Doney (University of Virginia)

Can phytoplankton help us determine ocean iron bioavailability?

Posted by mmaheigan

· Wednesday, March 11th, 2020

Iron (Fe) is a key element to sustaining life, but it is present at extremely low concentrations in seawater. This scarcity limits phytoplankton growth in large swaths of the global ocean, with implications for marine food webs and carbon cycling. The acquisition of Fe by phytoplankton is an important process that mediates the movement of carbon to the deep ocean and across trophic levels. It is a challenge to evaluate the ability of marine phytoplankton to obtain Fe from seawater since it is bound by a variety of poorly defined organic complexes.

Figure 1: Schematic representation of the reactions governing dissolved Fe (dFe) bioavailability to phytoplankton (a) Bioavailability of dFe in seawater collected from various basins and depth and probed with different iron-limited phytoplankton species under dim laboratory light and sunlight (b) (See paper for further details on samples and species)

A recent study in The ISME Journal proposes a new approach for evaluating seawater dissolved Fe (dFe) bioavailability based on its uptake rate constant by Fe-limited cultured phytoplankton. The authors collected samples from distinct regions across the global ocean, measured the properties of organic complexation, loaded these complexes with a radioactive Fe isotope, and then tracked the internalization rates from these forms to a diverse set of Fe-limited phytoplankton species. Regardless of origin, all of the phytoplankton acquired natural organic complexes at similar rates (accounting for cell surface area). This confirms that multiple Fe-limited phytoplankton species can be used to probe dFe bioavailability in seawater. Among water types, dFe bioavailability varied by ~4-fold and did not clearly correlate with Fe concentrations or any of the measured Fe speciation parameters. This new approach provides a novel way to determine Fe bioavailability in samples from across the oceans and enables modeling of in situ Fe uptake rates by phytoplankton based simply on measured Fe concentrations.

Authors:
Yeala Shaked (Hebrew University of Jerusalem)
Kristen N. Buck (University of South Florida)
Travis Mellett (University of South Florida)
Maria. T. Maldonado (University of British Columbia)

Pumped up by the cold: Increased elemental density in polar diatoms

Posted by mmaheigan

· Monday, October 28th, 2019

Large diatoms are common in polar phytoplankton blooms, contributing significantly to food webs and carbon export, but relatively little is known about their elemental biogeochemistry. A recent study in Frontiers in Marine Science showed that the size-dependent increase in cell nutrient content for polar diatoms was similar to published values for temperate diatoms, whereas the elemental density (mass per unit volume) of polar diatoms was substantially greater for all elements measured (carbon, nitrogen, silicon and phosphorus). Furthermore, at near freezing culture temperatures, there was a positive relationship between diatom size and realized growth rates near their theoretical maximum (Figure 1). Because of the differences in elemental density between carbon and silica, these diatoms exhibited particulate C:Si ratios that are commonly interpreted as a sign of iron limitation; yet these cultures were trace metal-replete. The observed elemental composition differences suggest that it may be important for polar biogeochemical models to include different representations of diatom biogeochemistry by accounting for the functions of size and near freezing temperature.

Figure 1. Left: Cellular carbon content for polar diatoms across four orders of magnitude in biovolume compared to the same relationship for a wide range of non-polar diatoms (MD&L = Menden-Deuer & Lessard, 2000). The y-intercept is the estimate of the baseline carbon density in these polar diatoms, and is significantly higher than the literature values reviewed in MD&L (2000). Right: Growth rate of the same polar diatoms expressed as a percent of their calculated maximum growth rate at 2°C. Error bars represent the range of values observed in the experiments. Maximum growth rate was estimated by 1) applying the growth rate/biovolume relationships published by Chisholm (1992) and Edwards et al. (2012) to the observed biovolume for each culture, and 2) scaling this growth rate to 2°C growth temperature using the relationship of Eppley (1972).

Authors:
Michael Lomas (Bigelow Laboratory for Ocean Sciences)
Steven Baer (Maine Maritime Academy)
Sydney Acton (Dauphin Island Sea Lab)
Jeffrey Krause (Dauphin Island Sea Lab and University of South Alabama)

The ecology of the biological carbon pump

Posted by mmaheigan

· Tuesday, October 15th, 2019

Plankton in the surface ocean convert CO₂ into organic biomass thereby fueling marine food webs. Part of this organic biomass sinks down into the deep ocean, where the surface-derived organic carbon, or respired CO₂, is locked in for decades to millennia. Without the biological carbon pump, atmospheric CO₂ would be ~200 ppm higher than it is today. We know that ecological processes in the surface ocean plankton communities have a paramount importance on the efficiency of the biological carbon pump. Unfortunately, however, the mechanisms how ecology determines sinking fluxes are poorly understood.

A recent study in Global Biogeochemical Cycles used large-scale in situ mesocosms to explore how the ecological interplay within plankton communities affects the downward flux of organic material. Organic biomass tends to sink faster when produced by smaller organisms because the sinking material they generate forms dense aggregates. Conversely, larger organisms produce relatively porous particles that sink more slowly.

Figure: Flow chart illustrating how plankton community structure affects the properties of sinking organic particles and ultimately the strength and efficiency of the biological carbon pump. The thick arrows at the bottom indicate that flux attenuation depends on the properties of particulate matter formed in the surface ocean. For example, slow-sinking porous aggregates containing large amounts of easily degradable organic substances will decay faster (right side) than dense aggregates of more refractory organic matter (left side).

The key finding of this study was the unexpectedly large influence that plankton community composition has on the degradation rate of sinking organic biomass. In fact, degradation rates changed maximally 15-fold over the course of the study while sinking speed changed only 3-fold. Degradation rate of sinking material, measured in oxygen consumption assays, was quite variable and tended to be higher for more easily degradable fresh organic matter. The rate was lower during harmful algal blooms, which produce toxic substances that inhibit organisms that feed on aggregates thereby reducing degradation rates. These findings are an important step forward as they show that our predictive understanding of the biological carbon pump could be improved substantially when linking degradation rates of sinking material with ecological processes in surface ocean plankton communities.

Authors:
L. T. Bach (University of Tasmania)
P. Stange, J. Taucher, E. P. Achterberg, M. Esposito, U. Riebesell (GEOMAR)
M. Algueró‐Muñiz (Alfred-Wegener-Institut Helmholtz)
H. Horn (NIOZ and Utrecht University)

Where the primary production goes determines whether you catch tuna or cod

Posted by mmaheigan

· Friday, September 6th, 2019

Fishes are incredibly diverse, fill various roles in their ecosystems, and are an important resource—economically, socially, and nutritionally. The relationship between primary productivity and fish catches is not straightforward; fisheries oceanographers and managers have long struggled to predict abundances and fully understand the controls of cross-ecosystem differences in fish abundances and assemblages. A recent study in Progress in Oceanography modeled the relationships between fish abundances and assemblages and ecosystem factors such as physical properties and plankton productivity.

The mechanistic model simulated feeding, growth, reproduction, and mortality of small pelagic forage fish, large pelagic fish, and demersal (bottom-dwelling) fish in the global ocean using plankton food web estimates and ocean conditions from a high-resolution earth system model of the 1990s. Modeled fish assemblages were more related to the separation of secondary production into pelagic zooplankton or benthic fauna secondary production than to primary productivity. Specifically, the ratio of pelagic to benthic production drove spatial differences in dominance by large pelagic fish or by demersal fish. Similarly, demersal fish abundance was highly sensitive to the efficiency of energy transfer from exported surface production to benthic fauna.

The model results offer a systematic understanding of how marine fish communities are structured by spatially varying environmental conditions. With global climate change, the expected decrease in exported primary production would lead to fewer demersal fish around the world. This model provides a framework for testing the effect of changing conditions on fish communities at a global scale, which can also help inform managers of potential impacts on economic, social, and nutritional resources worldwide.

Figure 1: (A) Sample food web with three fish types, two habitats, two prey categories, and feeding interactions (arrows). Dashed arrow denotes feeding only occurs in shelf regions with depth <200 m. (B) Fraction of large pelagic vs. demersal fishes (LP/(LP+D)) as a function of the ratio of zooplankton production lost to higher predation (Zoop) to detritus flux to the seafloor (Bent) averaged over large marine ecosystems. Solid line: predicted linear model response, dashed lines: standard error. (Lower panels) Circles=mean biomasses (g m^-2) and lines=fluxes of biomass (g m^-2 d^-1) through the pelagic (top 100m) and benthic components of the food webs at two test locations, (C) Peruvian Upwelling (PUP) ecosystem and (D) Eastern Bering Sea (EBS) shelf ecosystem. Circles and lines scale with the modeled biomasses and fluxes. Circle color key: Gray=net primary productivity (NPP); yellow=medium and large zooplankton; red=forage fish; blue=large pelagic fish; brown=benthos; green=demersal fish.

Authors:
Colleen M. Petrik (Princeton University, Texas A&M University)
Charles A. Stock (NOAA Geophysical Fluid Dynamics Laboratory)
Ken H. Andersen (Technical University of Denmark)
P. Daniël van Denderen (Technical University of Denmark)
James R. Watson (Oregon State University)

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.

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