Combined effects of different CO2 levels and N sources on the diazotrophic cyanobacterium Trichodesmium

To predict effects of climate change and possible feedbacks, it is crucial to understand the mechanisms behind CO2 responses of biogeochemically relevant phytoplankton species. Previous experiments on the abundant N2 fixers Trichodesmium demonstrated strong CO2 responses, which were attributed to an energy reallocation between its carbon (C) and nitrogen (N) acquisition. Pursuing this hypothesis, we manipulated the cellular energy budget by growing Trichodesmium erythraeum IMS101 under different CO2 partial pressure (pCO2) levels (180, 380, 980 and 1400?µatm) and N sources (N2 and NO3-). Subsequently, biomass production and the main energy-generating processes (photosynthesis and respiration) and energy-consuming processes (N2 fixation and C acquisition) were measured. While oxygen fluxes and chlorophyll fluorescence indicated that energy generation and its diurnal cycle was neither affected by pCO2 nor N source, cells differed in production rates and composition. Elevated pCO2 increased N2 fixation and organic C and N contents. The degree of stimulation was higher for nitrogenase activity than for cell contents, indicating a pCO2 effect on the transfer efficiency from N2 to biomass. pCO2-dependent changes in the diurnal cycle of N2 fixation correlated well with C affinities, confirming the interactions between N and C acquisition. Regarding effects of the N source, production rates were enhanced in NO3-grown cells, which we attribute to the higher N retention and lower ATP demand compared with N2 fixation. pCO2 effects on C affinity were less pronounced in NO3- users than N2 fixers. Our study illustrates the necessity to understand energy budgets and fluxes under different environmental conditions for explaining indirect effects of rising pCO2.

Seawater carbonate chemistry and processes during experiments with cyanobacterium Trichodesmium (IMS101), 2009

We investigated carbon acquisition by the N2-fixing cyanobacterium Trichodesmium IMS101 in response to CO2 levels of 15.1, 37.5, and 101.3 Pa (equivalent to 150, 370, and 1000 ppm). In these acclimations, growth rates as well as cellular C and N contents were measured. In vivo activities of carbonic anhydrase (CA), photosynthetic O2 evolution, and CO2 and HCO3- fluxes were measured using membrane inlet mass spectrometry and the 14C disequilibrium technique. While no differences in growth rates were observed, elevated CO2 levels caused higher C and N quotas and stimulated photosynthesis and N2 fixation. Minimal extracellular CA (eCA) activity was observed, indicating a minor role in carbon acquisition. Rates of CO2 uptake were small relative to total inorganic carbon (Ci) fixation, whereas HCO{3 contributed more than 90% and varied only slightly over the light period and between CO2 treatments. The low eCA activity and preference for HCO3- were verified by the 14C disequilibrium technique. Regarding apparent affinities, half-saturation concentrations (K1/2) for photosynthetic O2 evolution and HCO3- uptake changed markedly over the day and with CO2 concentration. Leakage (CO2 efflux : Ci uptake) showed pronounced diurnal changes. Our findings do not support a direct CO2 effect on the carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase (RubisCO) but point to a shift in resource allocation among photosynthesis, carbon acquisition, and N2 fixation under elevated CO2 levels. The observed increase in photosynthesis and N2fixation could have potential biogeochemical implications, as it may stimulate productivity in N-limited oligotrophic regions and thus provide a negative feedback in rising atmospheric CO2 levels.

Lucky Strike records of hydrothermal outflow temperatures

We deployed autonomous temperature sensors at black smoker chimneys, cracks, and diffuse flow areas at the Lucky Strike hydrothermal field (Mid-Atlantic Ridge, ~37°17'N) between summer 2009 and summer 2012 and contemporaneously measured tidal pressures and currents as part of the long-term MoMAR experiment to monitor hydrothermal activity. We classify the temperature data according to the hydrogeologic setting of the measurement sites: a high-temperature regime (>190°C) representing discharge of essentially unmixed, primary hydrothermal fluids through chimneys, an intermediate-temperature regime (10-100°C) associated with mixing of primary fluids with cold pore fluids discharging through cracks, and a low-temperature regime (<10°C) associated with a thermal boundary layer forming over bacterial mats associated with diffuse outflow of warm fluids. Temperature records from all the regimes exhibit variations at semi-diurnal tidal periods, and cross-spectral analyses reveal that high-temperature discharge correlates to tidal pressure while low-temperature discharge correlates to tidal currents. Intermediate-temperature discharge exhibits a transitional behavior correlating to both tidal pressure and currents. Episodic perturbations, with transient temperature drops of up to ~150°C, which occur in the high-temperature and intermediate-temperature records, are not observed on multiple probes (including nearby probes at the same site), and they are not correlated with microearthquake activity, indicating that the perturbation mechanism is highly localized at the measurement sites within the hydrothermal structures. The average temperature at a given site may increase or decrease at annual time scales, but the average temperature of the hydrothermal field, as a whole, appears to be stable over our 3 year observation period.

Field measurements of the atmosphere, ocean, sea ice and sub-ice platelet layer at Atka Bay in 2013

Ice shelves strongly interact with coastal Antarctic sea ice and the associated ecosystem by creating conditions favourable to the formation of a sub-ice platelet layer. The close investigation of this phenomenon and its seasonal evolution remain a challenge due to logistical constraints and a lack of suitable methodology. In this study, we characterize the seasonal cycle of Antarctic fast ice adjacent to the Ekström Ice Shelf in the eastern Weddell Sea. We used a thermistor chain with the additional ability to record the temperature response induced by cyclic heating of resistors embedded in the chain. Vertical sea-ice temperature and heating profiles obtained daily between November 2012 and February 2014 were analyzed to determine sea-ice and snow evolution, and to calculate the basal energy budget. The residual heat flux translated into an ice-volume fraction in the platelet layer of 0.18 ± 0.09, which we reproduced by a independent model simulation and agrees with earlier results. Manual drillings revealed an average annual platelet-layer thickness increase of at least 4m, and an annual maximum thickness of 10m beneath second-year sea ice. The oceanic contribution dominated the total sea-ice production during the study, effectively accounting for up to 70% of second-year sea-ice growth. In summer, an oceanic heat flux of 21 W/m**2 led to a partial thinning of the platelet layer. Our results further show that the active heating method, in contrast to the acoustic sounding approach, is well suited to derive the fast-ice mass balance in regions influenced by ocean/ice-shelf interaction, as it allows sub-diurnal monitoring of the platelet-layer thickness.

Zooplankton distribution and migration in low-oxygen modewater eddies

The eastern tropical North Atlantic (ETNA) features a mesopelagic oxygen minimum zone (OMZ) at approximately 300-600 m depth. Here, oxygen concentrations rarely fall below 40 µmol O2 kg-1, but are expected to decline under future projections of global warming. The recent discovery of mesoscale eddies that harbour a shallow suboxic (<5 µmol O2 kg-1) OMZ just below the mixed layer could serve to identify zooplankton groups that may be negatively or positively affected by on-going ocean deoxygenation. In spring 2014, a detailed survey of a suboxic anticyclonic modewater eddy (ACME) was carried out near the Cape Verde Ocean Observatory (CVOO), combining acoustic and optical profiling methods with stratified multinet hauls and hydrography. The multinet data revealed that the eddy was characterized by an approximately 1.5-fold increase in total area-integrated zooplankton abundance. At nighttime, when a large proportion of acoustic scatterers is ascending into the upper 150 m, a drastic reduction in mean volume backscattering (Sv, shipboard ADCP, 75kHz) within the shallow OMZ of the eddy was evident compared to the nighttime distribution outside the eddy. Acoustic scatterers were avoiding the depth range between about 85 to 120 m, where oxygen concentrations were lower than approximately 20 µmol O2 kg-1, indicating habitat compression to the oxygenated surface layer. This observation is confirmed by time-series observations of a moored ADCP (upward looking, 300kHz) during an ACME transit at the CVOO mooring in 2010. Nevertheless, part of the diurnal vertical migration (DVM) from the surface layer to the mesopelagic continued through the shallow OMZ. Based upon vertically stratified multinet hauls, Underwater Vision Profiler (UVP5) and ADCP data, four strategies have been identified to be followed by zooplankton in response to the eddy OMZ: i) shallow OMZ avoidance and compression at the surface (e.g. most calanoid copepods, euphausiids), ii) migration to the shallow OMZ core during daytime, but paying O2 debt at the surface at nighttime (e.g. siphonophores, Oncaea spp., eucalanoid copepods), iii) residing in the shallow OMZ day and night (e.g. ostracods, polychaetes), and iv) DVM through the shallow OMZ from deeper oxygenated depths to the surface and back. For strategy i), ii) and iv), compression of the habitable volume in the surface may increase prey-predator encounter rates, rendering zooplankton and micronekton more vulnerable to predation and potentially making the eddy surface a foraging hotspot for higher trophic levels. With respect to long-term effects of ocean deoxygenation, we expect avoidance of the mesopelagic OMZ to set in if oxygen levels decline below approximately 20 µmol O2 kg-1. This may result in a positive feedback on the OMZ oxygen consumption rates, since zooplankton and micronekton respiration within the OMZ as well as active flux of dissolved and particulate organic matter into the OMZ will decline.

Mesozooplankton abundance data from Disko Bay, West Greenland, 2008

The study site was located in the Disko Bay off Qeqertarsuaq, western Greenland. Due to land-connected sea ice coverage during winter, 2 sampling sites were combined. At the first site in winter (21 February to 23 March 2008), sampling was conducted through a hole in the ice at ca. 65 to 160 m depth approximately 0.5 nautical mile (n mile) south of Qeqertarsuaq (69° 14' N, 53° 29' W). In spring and summer (9 April to 18 July), sampling was done at a monitoring station 1 n mile south from Qeqertarsuaq (69° 14' N, 53° 23' W) at 300 m depth. Sampling was carried out between 10:00 and 17:00 h. During sampling from the ice, mesozooplankton was collected using a modified WP-2 net (45 µm) equipped with a closing mechanism (Hydrobios). Samples were collected in 3 depth strata (0-50, 50-100, and 100-150 m). During ship-based sampling, mesozooplankton was collected with a multinet (50 µm) equipped with a flow meter (Multinet, Hydrobios type midi), and 2 additional depth strata (150-200m and 200-250 m) were included. In addition to the seasonal study one diurnal investigation with sampling every 6 h was conducted from 29 April at 12:00 h to 30 April 30 at 12:00 h. Samples were immediately preserved in buffered formalin (5% final concentration) for later analyses. Biomass values of the different copepod species were calculated based on measurements of prosome length, and length/weight relationships. Two regressions for Calanus spp. were established for biomass calculations: one applicable prior to and during the phytoplankton bloom until 4 May, and another from 9 May onwards.