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Size does matter: larger krill leads to more POC export in the West Antarctic Peninsula

Posted by mmaheigan

· Friday, December 1st, 2023

Despite the importance of particulate organic carbon (POC) export on carbon sequestration and marine ecology, there have been few multi-decade studies in the world’s oceans. A new analysis published in Nature analyzed two decades of POC export data in the West Antarctic Peninsula and found that export oscillates on a 5-year cycle.

Figure caption: A) Particulate organic carbon (POC) export oscillates on a 5-year timescale in sync with the oscillation in the body size of the krill Euphausia superba on the West Antarctic Peninsula. B) POC export is significantly correlated with krill body size (p = 0.01).

Using a unique combination of krill data from penguin diet samples and net tows over two decades, Trinh et al. found that the cycle of POC export is intimately tied to the Antarctic krill (Euphausia superba) life cycle, as the bulk of the POC in their sediment traps was krill fecal pellets. Surprisingly, more krill did not lead to more POC export. Instead, when the krill population size was smaller but dominated by larger, older adults, POC export increased.

E. superba is the longest-lived (5-6 years) and largest krill species. They exhibit continuous annual growth throughout their life cycle. After about five years a krill population reaches its end stage and the population size is at a minimum. This end-stage population is composed of large, 50-60 mm long individuals that produce large, fast-sinking fecal pellets, leading to increased POC export. Increasing temperatures and deterioration of sea ice cover during the winter season due to climate change will likely impact the recruitment of new cohorts of krill and their success in replenishing aging populations. It is unclear how changes in the krill population and life cycle will impact long-term carbon sequestration on the West Antarctic Peninsula and nutrients exported to the benthic ecosystem

Authors:
Rebecca Trinh (Lamont Doherty Earth Observatory, Columbia University)
Hugh Ducklow (Lamont Doherty Earth Observatory, Columbia University)
Deborah Steinberg (Virginia Institute of Marine Science, College of William and Mary)
William Fraser (Polar Oceans Research Group)

Air-sea gas disequilibrium drove deoxygenation of the deep ice-age ocean

Posted by mmaheigan

· Thursday, March 18th, 2021

During the Last Glacial Maximum (~20,000 years ago, LGM) sediment data show that the deep ocean had lower dissolved oxygen (O₂) concentrations than the preindustrial ocean, despite cooler temperatures of this period increasing O₂ solubility in sea water.

Figure 1. a) Whole ocean inventory of the O₂ components in the preindustrial control (PIC): total O₂ (O₂); the preformed components equilibrium O₂ (O₂ equilibrium), physical disequilibrium O₂ (O₂ diseq phys) and biologically-mediated disequilibrium (O₂ diseq bio); and O₂ respired from soft-tissue (O₂ soft). b) The difference in whole ocean inventory of O₂ components between the LGM and PIC simulations.

In a study published in Nature Geoscience, the authors provide one of the first explanations for glacial deoxygenation. The authors combined a data-constrained model of the preindustrial (PIC) and LGM ocean with a novel decomposition of O₂ to assess the processes affecting the oceanic distribution of oxygen. The decomposition allowed for the preformed disequilibrium O₂—the amount of oxygen that deviates from its solubility equilibrium value when at the surface—to be tracked, along with other contributions such as the O₂ consumed by bacterial respiration of organic matter. In the preindustrial ocean, a third of the subsurface oxygen deficit was a result of disequilibrium rather than oxygen consumed by bacteria. This contradicts previous assumptions (Figure 1a). Nearly 80% of the disequilibrium resulted from upwelling waters, depleted in O₂ due to respiration, not fully equilibrating before re-subduction into the ocean interior. This effect was even greater during the LGM (Figure 1b). The authors attributed this largely to the widespread presence of sea ice—which acts as a cap on the surface preventing the water from gaining oxygen from the atmosphere—in the ocean around Antarctica, with a smaller contribution from iron fertilization.

This study provides one of the first mechanistic explanations for LGM deep ocean deoxygenation. As the ocean is currently losing oxygen due to warming, the effect of other processes, including sea ice changes, could prove important for understanding long-term ocean oxygenation changes.

Authors
Ellen Cliff (University of Oxford)
Samar Khatiwala (University of Oxford)
Andreas Schmittner (Oregon State University)

Joint highlight with GEOTRACES International Project Office

Ice sheets mobilize trace elements for export downstream

Posted by mmaheigan

· Thursday, January 7th, 2021

Trace elements are essential micronutrients for life in the ocean and also serve as valuable fingerprints of chemical weathering. The behaviour of trace elements in the ocean has gained interest because some of these elements are found at vanishingly low concentrations that limit ecosystem productivity. Despite delivering >2000 km³ yr^-1 of freshwater to the polar oceans, ice sheets have largely been overlooked as major trace element sources. This is partly due to a lack of data on meltwater endmember chemistry beneath and emerging from the Greenland and Antarctic ice sheets, which cover 10% of Earth’s land surface area, and partly because meltwaters were previously assumed to be dilute compared to most river waters.

In a study published in PNAS, authors analysed the trace element composition of meltwaters from the Mercer Subglacial Lake, a hydrologically active subglacial lake >1000 m below the surface of the Antarctic Ice Sheet, and a meltwater river emerging from beneath a large outlet glacier of the Greenland Ice Sheet (Leverett Glacier). These subglacial meltwaters (i.e., water travelling along the ice-rock interface beneath an ice mass) contained much higher concentrations of trace elements than anticipated. For example, typically immobile elements like iron and aluminium were observed in the dissolved phase (<0.45 µm) at much higher concentrations than in mean river or open ocean waters (up to 20,900 nM for Fe and 69,100 nM for Al), but exhibited large size fractionation between colloidal/nanoparticulate (0.02 – 0.45 µm) and soluble (<0.02 µm) size fractions (Figure 1). Subglacial physical and biogeochemical weathering processes are thought to mobilize many of these trace elements from the bedrock and sediments beneath ice sheets and export them downstream. Antarctic subglacial meltwaters were more enriched in dissolved trace elements than Greenland Ice Sheet outflow, which is likely due to longer subglacial residence times, lack of dilution from surface meltwater inputs, and differences in underlying sediment geology.

These results indicate that ice sheet systems can mobilize large quantities of trace elements from the land to the ocean and serve as major contributors to regional elemental cycles (e.g., coastal Southern Ocean). In a warming climate with increasing ice sheet runoff, subglacial meltwaters will become an increasingly dynamic source of micronutrients to coastal oceanic ecosystems in the polar regions.

Figure caption: Leverett Glacier (Greenland Ice Sheet) and Mercer Subglacial Lake (Antarctic Ice Sheet) dissolved elemental concentrations (<0.45 µm) normalized to mean non-glacial riverine trace element concentrations (Gaillardet et al., 2014) and major element concentrations (Martin and Meybeck, 1979). Grey regions indicate ±50 % of the riverine mean. Although major elements can be significantly depleted compared to non-glacial rivers, trace elements are commonly similar to or enriched.

Authors:
Jon R. Hawkings (Florida State Univ and German Research Centre for Geosciences)
Mark L. Skidmore (Montana State Univ)
Jemma L. Wadham (Univ of Bristol, UK)
John C. Priscu (Montana State Univ)
Peter L. Morton (Florida State Univ)
Jade E. Hatton (Univ of Bristol, UK)
Christopher B. Gardner (Ohio State Univ)
Tyler J. Kohler (École Polytechnique Fédérale de Lausanne, Switzerland)
Marek Stibal (Charles University, Prague, Czech Republic)
Elizabeth A. Bagshaw (Cardiff Univ, UK)
August Steigmeyer (Montana State Univ)
Joel Barker (Univ of Minnesota)
John E. Dore (Montana State Univ)
W. Berry Lyons (Ohio State Univ)
Martyn Tranter (Univ of Bristol, UK)
Robert G. M. Spencer (Florida State Univ)
SALSA Science Team

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)

Profiling floats reveal fate of Southern Ocean phytoplankton stocks

Posted by mmaheigan

· Tuesday, September 1st, 2020

More observations are needed to constrain the relative roles of physical (advection), biogeochemical (downward export), and ecological (grazing and biological losses) processes in driving the fate of phytoplankton blooms in Southern Ocean waters. In a recent paper published in Nature Communications, authors used seven Biogeochemical Argo (BGC-Argo) floats that vertically profiled the upper ocean every ten days as they drifted for three years across the remote Sea Ice Zone of the Southern Ocean. Using the floats’ biogeochemical sensors (chlorophyll, nitrate, and backscattering) and regional ratios of nitrate consumption:chlorophyll synthesis, the authors developed a new approach to remotely estimate the fate of the phytoplankton stocks, enabling calculations of herbivory and of downward carbon export. The study revealed that the major fate of phytoplankton biomass in this region is grazing, which consumes ~90% of stocks. The remaining 10% is exported to depth. This pattern was consistent throughout the entire sea ice zone where the floats drifted, from 60°-69° South.

Figure Caption: Southern Ocean Chlorophyll a climatology and floats’ trajectories (top panel). Total losses of Chlorophyll a (including grazing and phytodetritus export, left panel). Phytodetritus export (right panel).

This study region comprises two of the three major krill growth and development areas—the eastern Weddell and King Haakon VII Seas and Prydz Bay and the Kerguelen Plateau—so the observed grazing was probably due to Antarctic krill, underscoring their pivotal importance in this ecosystem. Building upon the greater understanding of ocean ecosystems via satellite ocean colour development in the 1990s, BGC-Argo floats and this new approach will allow remote monitoring of the different fates of phytoplankton stocks and insights into the status of the ecosystem.

Authors:
Sebastien Moreau (Norwegian Polar Institute, Tromsø, Norway)
Philip Boyd (Institute for Marine and Antarctic Studies, Hobart, Australia)
Peter Strutton (Institute for Marine and Antarctic Studies, Hobart, Australia)

A close-up view of biomass controls in Southern Ocean eddies

Posted by mmaheigan

· Thursday, August 20th, 2020

Southern Ocean biological productivity is instrumental in regulating the global carbon cycle. Previous correlative studies associated widespread mesoscale activity with anomalous chlorophyll levels. However, eddies simultaneously modify both the physical and biogeochemical environments via several competing pathways, making it difficult to discern which mechanisms are responsible for the observed biological anomalies within them. Two recently published papers track Southern Ocean eddies in a global, eddy-resolving, 3-D ocean simulation. By closely examining eddy-induced perturbations to phytoplankton populations, the authors are able to explicitly link eddies to co-located biological anomalies through an underlying mechanistic framework.

Figure caption: Simulated Southern Ocean eddies modify phytoplankton division rates in different directions of depending on the polarity of the eddy and background seasonal conditions. During summer anticyclones (top right panel) deliver extra iron from depth via eddy-induced Ekman pumping and fuel faster phytoplankton division rates. During winter (bottom right panel) the extra iron supply is eclipsed by deeper mixed layer depths and elevated light limitation resulting in slower division rates. The opposite occurs in cyclones.

In the first paper, the authors observe that eddies primarily affect phytoplankton division rates by modifying the supply of iron via eddy-induced Ekman pumping. This results in elevated iron and faster phytoplankton division rates in anticyclones throughout most of the year. However, during deep mixing winter periods, exacerbated light stress driven by anomalously deep mixing in anticyclones can dominate elevated iron and drive division rates down. The opposite response occurs in cyclones.

The second paper tracks how eddy-modified division rates combine with eddy-modified loss rates and physical transport to produce anomalous biomass accumulation. The biomass anomaly is highly variable, but can exhibit an intense seasonal cycle, in which cyclones and anticyclones consistently modify biomass in different directions. This cycle is most apparent in the South Pacific sector of the Antarctic Circumpolar Current, a deep mixing region where the largest biomass anomalies are driven by biological mechanisms rather than lateral transport mechanisms such as eddy stirring or propagation.

It is important to remember that the correlation between chlorophyll and eddy activity observable from space can result from a variety of physical and biological mechanisms. Understanding the nuances of how these mechanisms change regionally and seasonally is integral in both scaling up local observations and parameterizing coarser, non-eddy resolving general circulation models with embedded biogeochemistry.

Authors:
Tyler Rohr (Australian Antarctic Partnership Program, previously at MIT/WHOI)
Cheryl Harrison (University of Texas Rio Grande Valley)
Matthew Long (National Center for Atmospheric Research)
Peter Gaube (University of Washington)
Scott Doney (University of Virginia)

Krillin’ it with poop: Highlighting the importance of Antarctic krill in ocean carbon and nutrient cycling

Posted by mmaheigan

· Tuesday, February 4th, 2020

Scientists have long known the role of Antarctic krill (Euphausia superba) in Southern Ocean ecosystems. Evidence is gathering about krill’s biogeochemical importance through releasing millions of faecal pellets in swarms and stimulating primary production through nutrient excretion. Here, we explore and synthesise the known impacts that this highly abundant and rather large species has on the environment. Krill exemplify how metazoa can play a dominant role in shaping ocean biogeochemistry, thus providing additional motivation for protecting certain harvested species.

Figure 1: The ecological roles of krill in Southern Ocean biogeochemical cycles, including releasing faecal pellets, excreting nutrients whilst grazing, and larval krill migrating throughout the water column, shedding exoskeletons, and feeding on the seabed.

A review published in Nature Communications uncovers at least 13 possible pathways by which Antarctic krill either influence the carbon sink or release fertilizing nutrients (Figure 1). Their large size (up to 7 cm) and swarming nature (millions of krill aggregate) enable krill to strongly impact ocean biogeochemistry. Swarms release large numbers of faecal pellets, overwhelming detritivores and resulting in a large sink of faecal carbon. Krill may physically mix nutrients from the deep ocean and become a decades-long carbon store in whale biomass. Antarctic krill larvae, which live near the sea-ice, undergo deeper diel vertical migrations compared to adult Antarctic krill (400 m vs. 200 m), so any carbon respired or faecal pellets released by larvae could remain in the deep ocean longer than those released by adult krill at a shallower depth; the larval krill contribution to carbon export has not been quantified. Furthermore, it is currently unknown how many krill larvae are removed from the Antarctic krill fishery as by-catch. Perhaps the biggest challenge in constraining the role of krill (adult and larvae) in biogeochemical cycles is our limited capacity to quantify the abundance and biomass of Antarctic krill, since shipboard sampling methods (nets or acoustics) have limited spatial and temporal coverage. Ultimately, the Southern Ocean is an important physical AND biological sink of carbon, and we must consider the role krill and other animals have in this cycle.

Figure 2: Processes in the biological carbon pump including the sinking of dead phytoplankton aggregates, zooplankton, krill and fish faecal pellets and dead animals. Microbial remineralisation is depicted through the return of particulate organic carbon to dissolved organic carbon (DOC) and eventually carbon dioxide.

Authors:
Emma Cavan (Imperial College London and University of Tasmania)
Anna Belcher (British Antarctic Survey)
Angus Atkinson (Plymouth Marine Laboratory)
Simeon Hill (British Antarctic Survey)
So Kawaguchi (Australian Antarctic Division)
Stacey McCormack (University of Tasmania)
Bettina Meyer (Alfred Wegener Institute for Polar and Marine Research and University of Oldenburg)
Stephen Nicol (University of Tasmania)
Lavenia Ratnarajah (University of Liverpool)
Katrin Schmidt (University of Plymouth)
Deborah Steinberg (Virginia Institute of Marine Science)
Geraint Tarling (British Antarctic Survey)
Philip Boyd (University of Tasmania and Antarctic Climate and Ecosystems Cooperative Research Centre)

Pteropod populations stable or increasing according to long-term study along the Western Antarctic Peninsula

Posted by mmaheigan

· Thursday, March 21st, 2019

Shelled pteropods (pelagic snails) are abundant planktonic predators and prey, linking grazers and higher trophic levels and contributing to the carbon cycle via consumption and excretion. Pteropods have been heralded as bioindicators of ocean acidification, given their aragonitic shell’s susceptibility to dissolution, which could ultimately lead to declining abundance. However, pteropod population dynamics are understudied, particularly in the Southern Ocean, a region predicted to be highly impacted by both warming and ocean acidification. In a recent publication in Limnology and Oceanography, long-term data sets from the Western Antarctic Peninsula show that while there is considerable interannual variability in pteropod abundance, populations have remained stable over the past 25 years, with some pteropod species (gymnosomes (non-shelled pteropod) overall, L. antarctica and C. pyramidata (shelled pteropods) regionally) even increasing during this period (Figure 1).

Figure 1. Annual pteropod abundance anomalies for the entire Palmer Antarctica Long-Term Ecological Research (LTER) study region along the Western Antarctic Peninsula. (a) Limacina helicina antarctica (shelled pteropod), (b) Gymnosomes – nonshelled pteropods that prey on shelled pteropods (p = 0.007, r² = 0.27), and (c) Clio pyramidata (shelled pteropod). Effect of environment on pteropod abundance. (d) SST vs. L. antarctica abundance, e) Sea ice advance vs. L. antarctica and Gymnosome abundance, (f) Sea ice retreat vs. C. pyramidata abundance. Data plotted are annual anomalies for each year of the time series (1993–2017). Sea ice advance is lagged 2-yr behind pteropod abundance (e.g., 2017 pteropod annual anomaly is plotted against 2015 sea ice advance annual anomaly) SST are lagged 1-yr behind L. antarctica abundance (e.g., 2017 L. antarctica annual anomaly is plotted against 2016 SST). Regression lines for significant linear relationships are shown, regression statistics are as follows: (d) SST vs. L. antarctica (circles): n = 25, p = 0.006, r² = 0.25 (e) sea ice advance vs. L. antarctica (filled-circles) and Gymnosomes (empty-circles): n = 25, p = 0.003, r² = 0.30 (dashed line); (f) sea ice retreat vs. C. pyramidata (squares): n = 14, p = 0.0003, r² = 0.64.

There was no significant influence of carbonate chemistry parameters (e.g., aragonite saturation state) on pteropod abundance, since the Western Antarctic Peninsula has yet to experience prolonged conditions characteristic of ocean acidification. However, other environmental factors such as warming and associated sea ice retreat were more influential. For example, warmer, ice-free waters in one year typically led to higher pteropod abundances the following year, suggesting that pteropods may be better adapted than expected to warming conditions due to climate change. The authors propose that earlier sea ice retreat promotes recruitment and subsequent expansion of pteropods further South, which could explain their increased abundance in this subregion. These results increase our understanding of pteropod responses to environmental variability, which is important for predicting future effects of climate change on regional carbon cycling and plankton trophic interactions in the Southern Ocean.

Authors:
Patricia S. Thibodeau (VIMS)
Deborah K. Steinberg (VIMS)
Sharon E. Stammerjohn (University of Colorado at Boulder)
Claudine Hauri (University of Alaska Fairbanks)

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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