Anja Engel

Changes in biogenic carbon flow in response to sea surface warming

The pelagic ocean harbors one of the largest ecosystems on Earth. It is responsible for approximately half of global primary production, sustains worldwide fisheries, and plays an important role in the global carbon cycle. Ocean warming caused by anthropogenic climate change is already starting to impact the marine biota, with possible consequences for ocean productivity and ecosystem services. Because temperature sensitivities of marine autotrophic and heterotrophic processes differ greatly, ocean warming is expected to cause major shifts in the flow of carbon and energy through the pelagic system. Attempts to integrate such biological responses into marine ecosystem and biogeochemical models suffer from a lack of empirical data. Here, we show, using an indoor-mesocosm approach, that rising temperature accelerates respiratory consumption of organic carbon relative to autotrophic production in a natural plankton community. Increasing temperature by 2–6 °C hence decreased the biological drawdown of dissolved inorganic carbon in the surface layer by up to 31%. Moreover, warming shifted the partitioning between particulate and dissolved organic carbon toward an enhanced accumulation of dissolved compounds. In line with these findings, the loss of organic carbon through sinking was significantly reduced at elevated temperatures. The observed changes in biogenic carbon flow have the potential to reduce the transfer of primary produced organic matter to higher trophic levels, weaken the oceans biological carbon pump, and hence provide a positive feedback to rising atmospheric CO2.

Shifts in biogenic carbon flow from particulate to dissolved forms under high carbon dioxide and warm ocean conditions

Photosynthesis by phytoplankton in sunlit surface waters transforms inorganic carbon and nutrients into organic matter, a portion of which is subsequently transported vertically through the water column by the process known as the biological carbon pump (BCP). The BCP sustains the steep vertical gradient in total dissolved carbon, thereby contributing to net carbon sequestration. Any changes in the vertical transportation of the organic matter as a result of future climate variations will directly affect surface ocean carbon dioxide (CO 2) concentrations, and subsequently influence oceanic uptake of atmospheric CO 2 and climate. Here we present results of experiments designed to investigate the potential effects of ocean acidification and warming on the BCP. These perturbation experiments were carried out in enclosures (3,000 L volume) in a controlled mesocosm facility that mimicked future pCO 2 (∼900 ppmv) and temperature (3°C higher than ambient) conditions. The elevated CO 2 and temperature treatments disproportionately enhanced the ratio of dissolved organic carbon (DOC) production to particulate organic carbon (POC) production, whereas the total organic carbon (TOC) production remained relatively constant under all conditions tested. A greater partitioning of organic carbon into the DOC pool indicated a shift in the organic carbon flow from the particulate to dissolved forms, which may affect the major pathways involved in organic carbon export and sequestration under future ocean conditions.

Comment on "Phytoplankton Calcification in a High-CO2 World"

Iglesias-Rodriguez et al. (Research Articles, 18 April 2008, p. 336) reported that the coccolithophore Emiliania huxleyi doubles its organic matter production and calcification in response to high carbon dioxide partial pressures, contrary to previous laboratory and field studies. We argue that shortcomings in their experimental protocol compromise the interpretation of their data and the resulting conclusions.

Stimulated Bacterial Growth under Elevated pCO2: Results from an Off-Shore Mesocosm Study

Marine bacteria are the main consumers of freshly produced organic matter. Many enzymatic processes involved in the bacterial digestion of organic compounds were shown to be pH sensitive in previous studies. Due to the continuous rise in atmospheric CO2 concentration, seawater pH is presently decreasing at a rate unprecedented during the last 300 million years but the consequences for microbial physiology, organic matter cycling and marine biogeochemistry are still unresolved. We studied the effects of elevated seawater pCO2 on a natural plankton community during a large-scale mesocosm study in a Norwegian fjord. Nine Kiel Off-Shore Mesocosms for Future Ocean Simulations (KOSMOS) were adjusted to different pCO2 levels ranging initially from ca. 280 to 3000 µatm and sampled every second day for 34 days. The first phytoplankton bloom developed around day 5. On day 14, inorganic nutrients were added to the enclosed, nutrient-poor waters to stimulate a second phytoplankton bloom, which occurred around day 20. Our results indicate that marine bacteria benefit directly and indirectly from decreasing seawater pH. During the first phytoplankton bloom, 5–10% more transparent exopolymer particles were formed in the high pCO2 mesocosms. Simultaneously, the efficiency of the protein-degrading enzyme leucine aminopeptidase increased with decreasing pH resulting in up to three times higher values in the highest pCO2/lowest pH mesocosm compared to the controls. In general, total and cell-specific aminopeptidase activities were elevated under low pH conditions. The combination of enhanced enzymatic hydrolysis of organic matter and increased availability of gel particles as substrate supported up to 28% higher bacterial abundance in the high pCO2 treatments. We conclude that ocean acidification has the potential to stimulate the bacterial community and facilitate the microbial recycling of freshly produced organic matter, thus strengthening the role of the microbial loop in the surface ocean.

Floating ice-algal aggregates below melting arctic sea ice.

During two consecutive cruises to the Eastern Central Arctic in late summer 2012, we observed floating algal aggregates in the melt-water layer below and between melting ice floes of first-year pack ice. The macroscopic (1-15 cm in diameter) aggregates had a mucous consistency and were dominated by typical ice-associated pennate diatoms embedded within the mucous matrix. Aggregates maintained buoyancy and accumulated just above a strong pycnocline that separated meltwater and seawater layers. We were able, for the first time, to obtain quantitative abundance and biomass estimates of these aggregates. Although their biomass and production on a square metre basis was small compared to ice-algal blooms, the floating ice-algal aggregates supported high levels of biological activity on the scale of the individual aggregate. In addition they constituted a food source for the ice-associated fauna as revealed by pigments indicative of zooplankton grazing, high abundance of naked ciliates, and ice amphipods associated with them. During the Arctic melt season, these floating aggregates likely play an important ecological role in an otherwise impoverished near-surface sea ice environment. Our findings provide important observations and measurements of a unique aggregate-based habitat during the 2012 record sea ice minimum year.

Nitrogen processes in coastal and marine ecosystems

Nature of the problem • Nitrogen (N) inputs from human activities have led to ecological deteriorations in large parts of the coastal oceans along European coastlines, including harmful algae blooms and anoxia. • Riverine N-loads are the most pronounced nitrogen sources to coasts and estuaries. Other signifi cant sources are nitrogen in atmospheric deposition and fi xation. Approaches • Th is chapter describes all major N-turnover processes which are important for the understanding of the complexity of marine nitrogen cycling, including information on biodiversity. • Linkages to other major elemental cycles like carbon, oxygen, phosphorus and silica are briefl y described in this chapter. • A tentative budget of all major sources and sinks of nitrogen integrated for global coasts is presented, indicating uncertainties where present, especially the N-loss capacity of ocean shelf sediments. • Finally, specifi c nitrogen problems in the European Regional Seas, including the Baltic Sea, Black Sea, North Sea, and Mediterranean Sea are described. Key fi ndings/state of knowledge • Today, human activity delivers several times more nitrogen to the coasts compared to the natural background of nitrogen delivery. Th e source of this is the land drained by the rivers. Th erefore, the major European estuaries (e.g. Rhine, Scheldt, Danube and the coastlines receiving the outfl ow), North Sea, Baltic Sea, and Black Sea as well as some parts of the Mediterranean coastlines are aff ected by excess nutrient inputs. • Biodiversity is reduced under high nutrient loadings and oxygen defi ciency. Th is process has led to changes in the nutrient recycling in sediments, because mature communities of benthic animals are lacking in disturbed coastal sediments. Th e recovery of communities may not be possible if high productivity and anoxia persist for longer time periods. Major uncertainties/challenges • Th e magnitude of nitrogen sources are not yet well constrained. Likewise the role of nutrient ratios (N:P:Si ratios) may be a critical variable in the understanding of the development of harmful algae blooms. • Whether only inorganic forms of nitrogen are important for productivity, or whether organic nitrogen is also important is not well understood and needs future attention. Recommendations • For the future it will be necessary to develop an adaptive transboundary management strategy for nitrogen reduction. Th e starting point for such regulation is located in the catchments of rivers and along their way to the coastal seas. • An overall reduction of nitrogen inputs into the environment is urgently necessary, especially in the case of diff use nitrogen inputs from agricultural activities.

Phytoplankton blooms at increasing levels of atmospheric carbon dioxide: experimental evidence for negative effects on prymnesiophytes and positive on small picoeukaryotes

Anthropogenic emissions of carbon dioxide (CO2) and the ongoing accumulation in the surface ocean together with concomitantly decreasing pH and calcium carbonate saturation states have the potential to impact phytoplankton community composition and therefore biogeochemical element cycling on a global scale. Here we report on a recent mesocosm CO2 perturbation study (Raunefjorden, Norway), with a focus on organic matter and phytoplankton dynamics. Cell numbers of three phytoplankton groups were particularly affected by increasing levels of seawater CO2 throughout the entire experiment, with the cyanobacterium Synechococcus and picoeukaryotes (prasinophytes) profiting, and the coccolithophore Emiliania huxleyi (prymnesiophyte) being negatively impacted. Combining these results with other phytoplankton community CO2 experiments into a data-set of global coverage suggests that, whenever CO2 effects are found, prymnesiophyte (especially coccolithophore) abundances are negatively affected, while the opposite holds true for small picoeukaryotes belonging to the class of prasinophytes, or the division of chlorophytes in general. Future reductions in calcium carbonate-producing coccolithophores, providing ballast which accelerates the sinking of particulate organic matter, together with increases in picoeukaryotes, an important component of the microbial loop in the euphotic zone, have the potential to impact marine export production, with feedbacks to Earths climate system.

Summertime plankton ecology in Fram Strait - a compilation of long- and short-term observations

Between Greenland and Spitsbergen, Fram Strait is a region where cold ice-covered Polar Water exits the Arctic Ocean with the East Greenland Current (EGC) and warm Atlantic Water enters the Arctic Ocean with the West Spitsbergen Current (WSC). In this compilation, we present two different data sets from plankton ecological observations in Fram Strait: (1) long-term measurements of satellite-derived (1998–2012) and in situ chlorophyll a (chl a) measurements (mainly summer cruises, 1991–2012) plus protist compositions (a station in WSC, eight summer cruises, 1998–2011); and (2) short-term measurements of a multidisciplinary approach that includes traditional plankton investigations, remote sensing, zooplankton, microbiological and molecular studies, and biogeochemical analyses carried out during two expeditions in June/July in the years 2010 and 2011. Both summer satellite-derived and in situ chl a concentrations showed slight trends towards higher values in the WSC since 1998 and 1991, respectively. In contrast, no trends were visible in the EGC. The protist composition in the WSC showed differences for the summer months: a dominance of diatoms was replaced by a dominance of Phaeocystis pouchetii and other small pico- and nanoplankton species. The observed differences in eastern Fram Strait were partially due to a warm anomaly in the WSC. Although changes associated with warmer water temperatures were observed, further long-term investigations are needed to distinguish between natural variability and climate change in Fram Strait. Results of two summer studies in 2010 and 2011 revealed the variability in plankton ecology in Fram Strait.

Do submesoscale frontal processes ventilate the oxygen minimum zone off Peru

The Peruvian upwelling system encompasses the most intense and shallowest oxygen minimum zone (OMZ) in the ocean. This system shows pronounced submesoscale activity like filaments and fronts. We carried out glider-based observations off Peru during austral summer 2013 to investigate whether submesoscale frontal processes ventilate the Peruvian OMZ. We present observational evidence for the subduction of highly oxygenated surface water in a submesoscale cold filament. The subduction event ventilates the oxycline but does not reach OMZ core waters. In a regional submesoscale-permitting model we study the pathways of newly upwelled water. About 50% of upwelled virtual floats are subducted below the mixed layer within 5 days emphasizing a hitherto unrecognized importance of subduction for the ventilation of the Peruvian oxycline.

Effects of ocean acidification on the biogenic composition of the sea‐surface microlayer: Results from a mesocosm study

The sea-surface microlayer (SML) is the oceans uppermost boundary to the atmosphere and in control of climate relevant processes like gas exchange and emission of marine primary organic aerosols (POA). The SML represents a complex surface film including organic components like polysaccharides, proteins, and marine gel particles, and harbors diverse microbial communities. Despite the potential relevance of the SML in ocean-atmosphere interactions, still little is known about its structural characteristics and sensitivity to a changing environment such as increased oceanic uptake of anthropogenic CO2. Here we report results of a large-scale mesocosm study, indicating that ocean acidification can affect the abundance and activity of microorganisms during phytoplankton blooms, resulting in changes in composition and dynamics of organic matter in the SML. Our results reveal a potential coupling between anthropogenic CO2 emissions and the biogenic properties of the SML, pointing to a hitherto disregarded feedback process between ocean and atmosphere under climate change.