Particulate and phytoplankton absorption during POLARSTERN cruises ANT-XXVI/3 and ANT-XXVIII/3

The euphotic depth (Zeu) is a key parameter in modelling primary production (PP) using satellite ocean colour. However, evaluations of satellite Zeu products are scarce. The objective of this paper is to investigate existing approaches and sensors to estimate Zeu from satellite and to evaluate how different Zeu products might affect the estimation of PP in the Southern Ocean (SO). Euphotic depth was derived from MODIS and SeaWiFS products of (i) surface chlorophyll-a (Zeu-Chla) and (ii) inherent optical properties (Zeu-IOP). They were compared with in situ measurements of Zeu from different regions of the SO. Both approaches and sensors are robust to retrieve Zeu, although the best results were obtained using the IOP approach and SeaWiFS data, with an average percentage of error (E) of 25.43% and mean absolute error (MAE) of 0.10 m (log scale). Nevertheless, differences in the spatial distribution of Zeu-Chla and Zeu-IOP for both sensors were found as large as 30% over specific regions. These differences were also observed in PP. On average, PP based on Zeu-Chla was 8% higher than PP based on Zeu-IOP, but it was up to 30% higher south of 60°S. Satellite phytoplankton absorption coefficients (aph) derived by the Quasi-Analytical Algorithm at different wavelengths were also validated and the results showed that MODIS aph are generally more robust than SeaWiFS. Thus, MODIS aph should be preferred in PP models based on aph in the SO. Further, we reinforce the importance of investigating the spatial differences between satellite products, which might not be detected by the validation with in situ measurements due to the insufficient amount and uneven distribution of the data.

Plankton biomass in the Mamala Basin (Oahu Island, Hawaii Islands) under antropogenic influence in August-September of 2002-2004

Anthropogenic impact on biomass of coastal plankton communities caused by submerged disposal of urban sewage waters (dumping) was studied. Observations were carried out in August-September of 2002-2004 in the Mamala Bay (Oahu Island, Hawaii Islands) using satellite and straight sea measurements. An analysis of variability of integral indicators of the water column determined on the basis of on-board measurements allowed us to divide them into two groups: elements most sensitive to pollution (heterotrophic bacteria (H-Bact), phototrophic cyanobacteria Synechococcus spp. (SYN), and chlorophyll a (CHLa)) and elements that manifested episodic positive dependence on inflow of polluted waters (heterotrophic unicellular eukaryotes, small unicellular algae, phototrophic green bacteria Prochlorococcus spp., as well as total biomass of microplankton). It was shown that submerged waste water disposal in the region of the diffuser of the dumping device led to insignificant (aver. 1.2-1.4 times) local increase in integral biomass of H-Bact, SYN, and in concentration of CHLa. Similar but sharper (aver. 1.5-2.1 times) increase in these parameters was found in water layers with maximal biomasses. Possible pathways of disposed waters (under the pycnocline, at its upper boundary, and in the entire mixed layer) were analyzed on the basis of studying vertical displacement of biomasses of H-Bact, SYN, and prochlorophytes. Possibility of using optical anomalies distinguished from satellite data as markers of anthropogenic eutrophication caused by dumping was confirmed. Application of such markers depends on water transparency and on shapes of curves of vertical distribution of autotrophic organisms.

Physicochemical parameters of water sample the four coastal lagoons at the Peninsula of Yucatan

Study sites. Samples of surface water were taken from 4 coastal lagoons on the Yucatan Peninsula in Mexico: Celestun (20° 45' N - 90° 22' W), Chelem (21° 15' N - 89° 45' W), Rosada Lagoon (21º 19' N - 89º 19' W), and Sabancuy Estuary (18° 58' N - 91° 12' W). The sampling was performed from august to October of 2011 (Chelem 08/24; Laguna Rosada 09/06; Celestún 09/28; Sabancuy 10/25). The sampling was random without replacement and 10 samples of surface water were collected along a transect parallel to the coastal axis. Samples were deposited in sterile plastic bottles and conserved in refrigeration at 4°C. All samples were processed within 24 hours after sampling. According to the Mexican laws and regulations no permissions are required to obtain water and sediment samples from open public areas. Analysis of environmental and physicochemical parameters. Determinations of the environmental parameters were performed with a Hach 5465000 model 156 multi-parameter measuring instrument. The Lorenzen method was used to determine chlorophyll-a (21) with 90% acetone and the concentration was calculated according to the formula: Chla= 27.63 (OD665o - OD665a)(VA)/VM x L Where, OD665o: absorbance at 665 nm before acidification; OD665a: absorbance at 665 nm after acidification; VA: volume (ml) of acetone for extraction; VM: volume (ml) of filtered water; L: length (cm) of the photometric cell. Determinations of the physicochemical parameters (silicates, phosphates, nitrates, nitrites and ammonia) were performed using the spectrophotometric techniques described and modified by Strickland and Parsons (1972).

Carbon uptake rate calculated for melt pond water samples during POLARSTERN cruise ARK-XXVII/3 (IceArc) in 2012

Net Primary Production was measured using the 14**C uptake method with minor modifications. Melt pond samples were spiked with 0.1µCi ml**-1 of 14**C labelled sodium bicarbonate (Moravek Biochemicals, Brea, USA) and distributed in 10 clear bottles (20 ml each). Subsequently they were incubated for 12 h at -1.3°C under different scalar irradiances (0-420 µmol photons m**-2 s**-1) measured with a spherical sensor (Spherical Micro Quantum Sensor US-SQS/L, Heinz Walz, Effeltrich, Germany). At the end of the incubation, samples were filtered onto 0.2 µm nitrocellulose filters and the particulate radioactive carbon uptake was determined by liquid scintillation counting using Filter count scintillation cocktail (Perkin Elmer, Waltham, USA). The carbon uptake values in the dark were subtracted from the carbon uptake values measured in the light incubations. Dissolved inorganic carbon (DIC) was measured for each sample using the flow injection system (Hall and Aller, 1992). The DIC concentration was taken into account to calculate the amount of labeled bicarbonate incorporated into the cell. Carbon fixation rates were normalized volumetrically and by chlorophyll a. Photosynthesis-irradiance curves (PI curves) were fitted using MATLAB® according to the equation proposed by Platt et al. (1980) including a photoinhibition parameter (beta) and providing the main photosynthetic parameters: maximum Chla normalized carbon fixation rate if there were no photoinhibition (Pb) and the initial slope of the saturation curve (alpha). The derived parameters: light intensity at which photosynthesis is maximal (Im), the carbon fixation rate at that maximal irradiance (Pbm) and the adaptation parameter or photoacclimation index (Ik) were calculated according to Platt et al. (1982).

Carbon uptake rate calculated for CTD water samples during POLARSTERN cruise ARK-XXVII/3 (IceArc) in 2012

Net Primary Production was measured using the 14**C uptake method with minor modifications. Seawater samples were spiked with 0.1µCi ml**-1 of 14**C labelled sodium bicarbonate (Moravek Biochemicals, Brea, USA) and distributed in 10 clear bottles (20 ml each). Subsequently they were incubated for 12 h at -1.3°C under different scalar irradiances (0-420 µmol photons m**-2 s**-1) measured with a spherical sensor (Spherical Micro Quantum Sensor US-SQS/L, Heinz Walz, Effeltrich, Germany). At the end of the incubation, samples were filtered onto 0.2 µm nitrocellulose filters and the particulate radioactive carbon uptake was determined by liquid scintillation counting using Filter count scintillation cocktail (Perkin Elmer, Waltham, USA). The carbon uptake values in the dark were subtracted from the carbon uptake values measured in the light incubations. Dissolved inorganic carbon (DIC) was measured for each sample using the flow injection system (Hall and Aller, 1992). The DIC concentration was taken into account to calculate the amount of labeled bicarbonate incorporated into the cell. Carbon fixation rates were normalized volumetrically and by chlorophyll a. Photosynthesis-irradiance curves (PI curves) were fitted using MATLAB® according to the equation proposed by Platt et al. (1980) including a photoinhibition parameter (beta) and providing the main photosynthetic parameters: maximum Chla normalized carbon fixation rate if there were no photoinhibition (Pb) and the initial slope of the saturation curve (alpha). The derived parameters: light intensity at which photosynthesis is maximal (Im), the carbon fixation rate at that maximal irradiance (Pbm) and the adaptation parameter or photoacclimation index (Ik) were calculated according to Platt et al. (1982).

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.

Carbon uptake rate calculated for sea-ice core samples during POLARSTERN cruise ARK-XXVII/3 (IceArc) in 2012

Net Primary Production was measured using the 14**C uptake method with minor modifications. Melted sea ice samples were spiked with 0.1µCi ml**-1 of 14**C labelled sodium bicarbonate (Moravek Biochemicals, Brea, USA) and distributed in 10 clear bottles (20 ml each). Subsequently they were incubated for 12 h at -1.3°C under different scalar irradiances (0-420 µmol photons m**-2 s**-1) measured with a spherical sensor (Spherical Micro Quantum Sensor US-SQS/L, Heinz Walz, Effeltrich, Germany). At the end of the incubation, samples were filtered onto 0.2 µm nitrocellulose filters and the particulate radioactive carbon uptake was determined by liquid scintillation counting using Filter count scintillation cocktail (Perkin Elmer, Waltham, USA). The carbon uptake values in the dark were subtracted from the carbon uptake values measured in the light incubations. Dissolved inorganic carbon (DIC) was measured for each sample using the flow injection system (Hall and Aller, 1992). The DIC concentration was taken into account to calculate the amount of labeled bicarbonate incorporated into the cell. Carbon fixation rates were normalized volumetrically and by chlorophyll a. Photosynthesis-irradiance curves (PI curves) were fitted using MATLAB® according to the equation proposed by Platt et al. (1980) including a photoinhibition parameter (beta) and providing the main photosynthetic parameters: maximum Chla normalized carbon fixation rate if there were no photoinhibition (Pb) and the initial slope of the saturation curve (alpha). The derived parameters: light intensity at which photosynthesis is maximal (Im), the carbon fixation rate at that maximal irradiance (Pbm) and the adaptation parameter or photoacclimation index (Ik) were calculated according to Platt et al. (1982).

Macroalgal responses to ocean acidification depend on nutrient and light levels

Ocean acidification may benefit algae that are able to capitalize on increased carbon availability for photosynthesis, but it is expected to have adverse effects on calcified algae through dissolution. Shifts in dominance between primary producers will have knock-on effects on marine ecosystems and will likely vary regionally, depending on factors such as irradiance (light vs. shade) and nutrient levels (oligotrophic vs. eutrophic). Thus experiments are needed to evaluate interactive effects of combined stressors in the field. In this study, we investigated the physiological responses of macroalgae near a CO2 seep in oligotrophic waters off Vulcano (Italy). The algae were incubated in situ at 0.2 m depth using a combination of three mean CO2 levels (500, 700-800 and 1200 µatm CO2), two light levels (100 and 70% of surface irradiance) and two nutrient levels of N, P, and K (enriched vs. non-enriched treatments) in the non-calcified macroalga Cystoseira compressa (Phaeophyceae, Fucales) and calcified Padina pavonica (Phaeophyceae, Dictyotales). A suite of biochemical assays and in vivo chlorophyll a fluorescence parameters showed that elevated CO2 levels benefitted both of these algae, although their responses varied depending on light and nutrient availability. In C. compressa, elevated CO2 treatments resulted in higher carbon content and antioxidant activity in shaded conditions both with and without nutrient enrichment-they had more Chla, phenols and fucoxanthin with nutrient enrichment and higher quantum yield (Fv/Fm) and photosynthetic efficiency (alpha ETR) without nutrient enrichment. In P. pavonica, elevated CO2 treatments had higher carbon content, Fv/Fm, alpha ETR, and Chla regardless of nutrient levels-they had higher concentrations of phenolic compounds in nutrient enriched, fully-lit conditions and more antioxidants in shaded, nutrient enriched conditions. Nitrogen content increased significantly in fertilized treatments, confirming that these algae were nutrient limited in this oligotrophic part of the Mediterranean. Our findings strengthen evidence that brown algae can be expected to proliferate as the oceans acidify where physicochemical conditions, such as nutrient levels and light, permit.

Seagrass ecosystem response to long-term high CO2 in a Mediterranean volcanic vent

We examined the long-term effect of naturally acidified water on a Cymodocea nodosa meadow growing at a shallow volcanic CO2 vent in Vulcano Island (Italy). Seagrass and adjacent unvegetated habitats growing at a low pH station (pH = 7.65 ± 0.02) were compared with corresponding habitats at a control station (pH = 8.01 ± 0.01). Density and biomass showed a clear decreasing trend at the low pH station and the below- to above-ground biomass ratio was more than 10 times lower compared to the control. C content and delta 13C of leaves and epiphytes were significantly lower at the low pH station. Photosynthetic activity of C. nodosa was stimulated by low pH as seen by the significant increase in Chla content of leaves, maximum electron transport rate and compensation irradiance. Seagrass community metabolism was intense at the low pH station, with significantly higher net community production, respiration and gross primary production than the control community, whereas metabolism of the unvegetated community did not differ between stations. Productivity was promoted by the low pH, but this was not translated into biomass, probably due to nutrient limitation, grazing or poor environmental conditions. The results indicate that seagrass response in naturally acidified conditions is dependable upon species and geochemical characteristics of the site and highlight the need for a better understanding of complex interactions in these environments.

Land based mesocosm experiment studying the simultaneous impact of warming and acidification on the planktonic food web of the Eastern Mediterranean

A land based mesocosm experiment focusing on the study of the simultaneous impact of warming and acidification on the planktonic food web of the Eastern Mediterranean took place in August-September 2013 at the mesocosm facilities of HCMR in Crete (CRETACOSMOS). Two different pCO2 (present day and predicted for year 2100) were applied in triplicate mesocosms of 3 m**3. This was tested in two different temperatures (ambient seawater T and ambient T plus 3°C). Twelve mesocosms in total were incubated in two large concrete tanks. Temperature was controlled by sophisticated, automated systems. A large variety of chemical, biological and biochemical variables were studied, including salinity, temperature, light and alkalinity measurements, inorganic and organic, particulate and dissolved, nutrient analyses, biological stock (Chla concentration, enumeration and community composition of microbial, phyto- and zooplankton organisms) and rate (primary, bacterial, viral production, copepod egg production, zooplankton grazing, N2 fixation, P uptake) measurements, bacterial DNA extraction and phytoplankton transcriptomics, calcifiers analyses. Twenty three scientists from 6 Institutes and 5 countries participated in this experiment.

Effects of ocean acidification on different life-cycle stages of the kelp Laminaria hyperborea (Phaeophyceae), link to supplementary data

Our objective for this study was to evaluate the influence of preindustrial and expected future atmospheric CO2 concentrations (280 µatm and 700 µatm pCO2, respectively) on different life-cycle stages of the kelp Laminaria hyperborea from Helgoland (Germany, North Sea). Zoospore germination, gametogenesis, vegetative growth, sorus formation and photosynthetic performance of vegetative and fertile tissue were examined. The contribution of external carbonic anhydrase (exCA) to C-supply for net-photosynthesis (net-PS) and the Chla- and phlorotannin content were investigated. Female gametogenesis and vegetative growth of sporophytes were significantly enhanced under the expected future pCO2. rETR(max) and net-PS of young vegetative sporophytes tended to increase performance at higher pCO2. The trend towards elevated net-PS vanished after inhibition of exCA. In vegetative sporophytes, phlorotannin content and Chla content were not significantly affected by pCO2.

Sediment, water and biomass characteristics along gradients in Kongsfjorden, Svalbard

In contrast to numerous studies on the biomass of marine microphytobenthos from temperate coastal ecosystems, little is known from polar regions. Therefore, microphytobenthos biomass was measured at several coastal sites in Arctic Kongsfjorden (Spitsbergen) during the polar summer (June-August 2006). On sandy sediments, chla varied between 8 and 200 mg/m**2 and was related to water depth, current/wave exposure and geographical location. Biomass was rather independent of abiotic parameters such as sediment properties, salinity, temperature or light availability. At three stations, sediments at water depths of 3-4, 10, 15, 20 and 30 m were investigated to evaluate the effect of light availability on microalgae. Significant differences in distribution patterns of biomass in relation to deeper waters >10 m were found. The productive periods were not as distinct as phytoplankton blooms. Only at 3-4 m water depth at all three stations were two- to threefold increases of biomass measured during the investigation period. Hydrodynamic conditions seemed to be the driving force for differences in sediment colonisation by benthic microalgae. In spite of the extreme Arctic environmental conditions for algal growth, microphytobenthos biomass was comparable to marine temperate waters.

Underway measurements of dissolved gases along POLARSTERN cruise track ANT-XXVII/2

We present measurements of pCO2, O2 concentration, biological oxygen saturation (Delta O2/Ar) and N2 saturation (Delta N2) in Southern Ocean surface waters during austral summer, 2010-2011. Phytoplankton biomass varied strongly across distinct hydrographic zones, with high chlorophyll a (Chla) concentrations in regions of frontal mixing and sea-ice melt. pCO2 and Delta O2 /Ar exhibited large spatial gradients (range 90 to 450 µatm and -10 to 60%, respectively) and co-varied strongly with Chla. However, the ratio of biological O2 accumulation to dissolved inorganic carbon (DIC) drawdown was significantly lower than expected from photosynthetic stoichiometry, reflecting the differential time-scales of O2 and CO2 air-sea equilibration. We measured significant oceanic CO2 uptake, with a mean air-sea flux (~ -20 mmol m-2 d-1) that significantly exceeded regional climatological values. N2 was mostly supersaturated in surface waters (mean Delta N2 of +2.5 %), while physical processes resulted in both supersaturation and undersaturation of mixed layer O2 (mean Delta O2phys = 2.1 %). Box model calculations were able to reproduce much of the spatial variability of Delta N2 and Delta O2phys along the cruise track, demonstrating significant effects of air-sea exchange processes (e.g. atmospheric pressure changes and bubble injection) and mixed layer entrainment on surface gas disequilibria. Net community production (NCP) derived from entrainment-corrected surface Delta O2 /Ar data, ranged from ~ -40 to > 300 mmol O2 m-2 d-1 and showed good coherence with independent NCP estimates based on seasonal mixed layer DIC deficits. Elevated NCP was observed in hydrographic frontal zones and regions of sea-ice melt with shallow mixed layer depths, reflecting the importance of mixing in controlling surface water light and nutrient availability.