Temperature tolerance of western Baltic Sea Fucus vesiculosus - growth, photosynthesis and survival

Fucus vesiculosus L. (Phaeophyceae) is the most abundant and hence ecologically most important primary producer, carbon sink and habitat provider in the western Baltic Sea. All F. vesiculosus L. specimens were collected on 23 April 2014 from a depth of 0.2-1 m in the non-tidal Kiel Fjord, western Baltic Sea (54°27'N; 10°12'E), where this species forms dense and almost monospecific stands on stones. After sampling the algal thalli were stored in a refrigerator box with water from the sampling site, transported to Bremerhaven and stored at 10 °C for one day in filtered seawater. Experiments were conducted with vegetative apical tips (6.7±0.5 cm length), the actively growing region of F. vesiculosus, which were randomly selected and cut from 144 different individuals prior to the experiments. These tips were acclimated to laboratory conditions for three days in filtered seawater at 10 °C before the start of the experiment. Furthermore, 30 additional vegetative apices were freeze-dried to document the initial biochemical status of F. vesiculosus in its native habitat. A temperature gradient was installed in a walk-in constant cooling chamber (15 °C) in nine water baths (5, 10, 15, 20, 24, 26, 27, 28 and 29 °C ± 0.1 °C) which were tempered by thermostats (5, 10 and 15 °C: Huber Variostat CC + Pilot ONE, Peter Huber Kältemaschinen GmbH, Offenburg, Germany; 20 and 28 °C: Haake DC3, Thermo Fisher Scientific Inc., Waltham, USA; 24, 26, 27 and 29 °C: Haake DC10). Every temperature treatment consisted of four 2 L glass beakers (n = 4). In each beaker four F. vesiculosus apices were grown in 2 µm-filtered North Sea water diluted with demineralized water in a ratio of 1:1 and enriched with nutrients after Provasoli (1968; 1/10 enrichment), leading to a salinity of about 15.6 which equaled habitat conditions. The algae were exposed to an irradiance of 130 µmol photons m-2 s-1 ±10 % (Powerstar HGI-TS 150 W, OSRAM GmbH, Bad Homburg, Germany) measured at the top of the beaker under a 16:8 h L:D cycle. The media in the beakers was changed every third or fourth day and aerated with artificial air containing 380 ppm CO2 (gas mixing device; HTK Hamburg GmbH, Hamburg, Germany). Before the experiment, the algae were acclimated to the final temperatures in steps of 5 °C for 2 days each, beginning at 10 °C. After 21 days exposure time, three out of four samples per replicate were freeze-dried for further biochemical analyses, and afterwards the thermostats were turned off to reduce the temperature to 16±0.4 °C for another 10 days permitting growth under post-culture conditions.

Temperature proxies of sediment cores from the Sea of Marmara

Reconstructing ocean temperature values is a major target in paleoceanography and climate research. However, most temperature proxies are organism-based and thus suffer from an "ecological bias". Multiproxy approaches can potentially overcome this bias but typically require more investment in time and resources, while being susceptible to errors induced by sample preparation steps necessary before analysis. Three lipid-based temperature proxies are widely used: UK'37 (based on long chain alkenones from phytoplanktonic haptophytes), TEX86 [based on glycerol dialkyl glycerol tetraethers (GDGTs) from pelagic archaea] and LDI (based on long chain diols from phytoplanktonic eustigmatophytes). So far, separate analytical methods, including gas chromatography (GC) and liquid chromatography (LC), have been used to determine these proxies. Here we present a sensitive method for determining all three in a single normal phase high performance LC-atmospheric pressure chemical ionization mass spectrometry (NP-HPLC-APCI-MS) analysis. Each of the long chain alkenones and long chain diols was separated and unambiguously identified from the accurate masses and characteristic fragmentation during multiple stage MS analysis (MS2). Comparison of conventional GC and HPLC-MS methods showed similar results for UK'37 and LDI, respectively, using diverse environmental samples and an Emiliania huxleyi culture. Including the three sea surface temperature (SST) proxies; the NP-HPLC-APCI-MS method in fact allows simultaneous determination of nine paleoenvironmental proxies. The extent to which the ecology of the source organisms (ecological bias) influences lipid composition and thereby the reconstructed temperature values was demonstrated by applying the new method to a sediment core from the Sea of Marmara, covering the last 21 kyr BP. Reconstructed SST values differed considerably between the proxies for the Last Glacial Maximum (LGM) and the period of Sapropel S1 formation at ca. 10 kyr BP, whereas the trends during the late Holocene were similar. Changes in the composition of alkenone-producing species at the transition from the LGM to the Bølling/Allerød (B/A) were inferred from unreasonably high UK'37-derived SST values (ca. 20 °C) during the LGM. We ascribe discrepancies between the reconstructed temperature records during S1 deposition to habitat change, e.g. a different depth due to changes in nutrient availability.

Mesozooplankton investigations in Bornholm Basin, Baltic Sea

Mesozooplankton production was estimated by using a new sampling technique and two alternative calculation methods. In essence, production estimates are based on significantly higher abundances. The contribution of juvenile stages to copepod and fish dynamics was generally low, so that the omission of juvenile stages in budgets will result in a small error. The situations reported in this study present a unique food web szenario, which in detail, however, was strongly dependent on methodology. Furthermore relations between trophic levels were considered with respect to vertical distribution.

(Table 1) Diving behaviour of male Adélie penguins (Pygoscelis adeliae) with and without corticosterone implants at Pointe Géologie

The amount of energy that organisms can allocate to self-maintenance and/or reproduction largely depends on their foraging strategies. Because of corticosterone (CORT) involvement in the control of energy metabolism, food intake and locomotor activity, recent studies have sought to demonstrate the role of this hormone in foraging decisions and performance. Moreover, considerable recent advances in animal-attached loggers now allow the study of behaviour in free-living animals. In order to assess the effects of CORT administration on the foraging behaviour of free-living Adelie Penguins Pygoscelis adeliae, we studied a group with CORT implants and a control group without CORT implants, by attaching time-depth recorders to the two groups and monitoring them throughout up to seven consecutive foraging trips during the guard stage (in Adelie Land, Antarctica). We found that foraging trips duration was similar between both groups. Dive durations, time spent at the bottom phase of dives, and the number of undulations per dive of CORT-implanted birds were all significantly higher than those of controls. However, CORT-implanted birds performed fewer dives overall (ca. 4,400) than controls (ca. 6,250) and spent many (13 and 6 times for penguins #3 and #4, respectively) long periods (>3 h) without diving. The low foraging effort and long resting periods support the view that CORT-implanted birds probably gained less energy than did the control birds. CORT treatment appears then to result in redirecting bird behaviour from costly activity (i.e. reproduction) to a behaviour promoting the preservation of energy reserves. Future studies are therefore needed to assess body condition and reproductive success of CORT-manipulated birds in parallel with the recording of their diving performances.

Body mass, flipper length, oxidative balance indices and telomere length of king penguin chicks (A. patagonicus) on Possession Island

One of the reasons for animals not to grow as fast as they potentially could is that fast growth has been shown to be associated with reduced lifespan. However, we are still lacking a clear description of the reality of growth-dependent modulation of ageing mechanisms in wild animals. Using the particular growth trajectory of small king penguin chicks naturally exhibiting higher-than-normal growth rate to compensate for the winter break, we tested whether oxidative stress and telomere shortening are related to growth trajectories. Plasma antioxidant defences, oxidative damage levels and telomere length were measured at the beginning and at the end of the post-winter growth period in three groups of chicks (small chicks, which either passed away or survived the growth period, and large chicks). Small chicks that died early during the growth period had the highest level of oxidative damage and the shortest telomere lengths prior to death. Here, we show that small chicks that grew faster did it at the detriment of body maintenance mechanisms as shown by (i) higher oxidative damage and (ii) accelerated telomere loss. Our study provides the first evidence for a mechanistic link between growth and ageing rates under natural conditions.

Diversity of planktonic foraminifer fauna found in 18 deep-sea cores from the North Atlantic

Greenland stadial/interstadial cycles are known to affect the North Atlantic's hydrography and overturning circulation and to cause ecological changes on land (e.g., vegetation). Hardly any information, directly expressed as diversity indices, however, exists on the impacts of these millennial-scale variations on the marine flora and fauna. We calculated three diversity indices (species richness, Shannon diversity index, Hurlbert's probability of interspecific encounter) for the planktonic foraminifer fauna found in 18 deep-sea cores covering a time span back to 60 ka. Clear differences in diversity response to the abrupt climate change can be observed and some records can be grouped accordingly. Core SO82-05 from the southern section of the subpolar gyre, the cores along the British margin and core MD04-2845 in the Bay of Biscay show two modes of diversity distribution, with reduced diversity (uneven fauna) during cold phases and the reverse (even fauna) during warm phases. Along the Iberian margin high species diversity prevailed throughout most of the glacial period. The exceptions were the Heinrich stadials when the fauna abruptly shifted from an even to an uneven or less even fauna. Diversity changes were often abrupt, but revealed a high resilience of the planktonic foraminifer faunas. The subtropical gyre waters seem to buffer the climatic effects of the Heinrich events and Greenland Stadials allowing for a quick recovery of the fauna after such an event. The current work clearly shows that planktonic foraminifer faunas quickly adapt to climate change, albeit with a reduced diversity.

Copepod abundance and fecal pellet production, sinking rate, and degradation in samples obtained during Leg 7 of Galathea 3 cruise 2006

The vertical distribution of copepods, fecal pellets and the fecal pellet production of copepods were measured at seven stations across the Southern Indian Ocean from productive areas off South Africa to oligotrophic waters off Northern Australia during October/November 2006. We quantified export of copepod fecal pellet from surface waters and how much was retained. Furthermore, the potential impact of Oncaea spp. and harpacticoid copepods on fecal pellets degradation was evaluated and found to be regional substantial. The highest copepod abundance and fecal pellet production was found in the western nutrient-rich stations close to South Africa and the lowest at the central oligotrophic stations. The in situ copepod fecal pellet production varied between 1 and 1,000 µg C/m**3/day. At all stations, the retention of fecal pellets in the upper 400 m of the water column was more than 99% and the vertical export of fecal pellets was low (<0.02 mg/m**2/day).

Organic geochemistry records from two Palaeocene-Eocene Thermal Maximum sections in Denmark

Analysis of sediments deposited at different latitudes around the world during the Palaeocene-Eocene Thermal Maximum (PETM; ~56 Ma) have revealed a globally profound warming phase, regionally varying from 5-8 °C. Such records from Europe have not yet been obtained. We studied the variations in sea surface and continental mean annual air temperatures (SST and MAT, respectively) and the distribution patterns and stable carbon isotopes of higher plant derived n-alkanes in two proximal PETM sections (Fur and Store Bælt, Denmark) from the epicontinental North Sea Basin. A negative carbon isotope excursion (CIE) of 4-7 per mil was recorded in land plant derived n-alkanes, similar to what has been observed for other PETM sections. However, differences observed between the two proximal sites suggest that local factors, such as regional vegetation and precipitation patterns, also influenced the CIE. The presence of S-bound isorenieratene derivatives at the onset of the PETM and increased organic carbon contents points to a rapid shift in depositional environment; from well oxygenated to anoxic and sulfidic. These euxinic conditions are comparable with those during the PETM in the Arctic Ocean. SSTs inferred from TEX86 show relatively low temperatures followed by an increase of ~7 °C across the PETM. At the Fur section, a remarkably similar temperature record was obtained for MAT using the MBT'/CBT proxy. However, the MAT record of the Store Bælt section did not reveal this warming.

Impact of ocean acidification and warming on respiration, heart rate and acid-base status of Mytilus edulis from the North Sea

Anthropogenic climate change confronts marine organisms with rapid trends of concomitant warming and CO2 induced ocean acidification. The survival and distribution of species partly depend on their ability to exploit their physiological plasticity during acclimatization. Therefore, in laboratory studies the effects of simulated future ocean acidification on thermal tolerance, energy metabolism and acid-base regulation capacity of the North Sea population of the blue mussel Mytilus edulis were examined. Following one month of pre-acclimation to 10 °C and control CO2 levels, mussels were exposed for two weeks to control and projected oceanic CO2 levels (390, 750 and 1120 µatm) before being subjected to a stepwise warming protocol between 10 °C and 31 °C (+ 3 °C each night). Oxygen consumption and heart rates, anaerobic metabolite levels and haemolymph acid-base status were determined at each temperature. CO2 exposure left oxygen consumption rate unchanged at acclimation temperature but caused a somewhat stronger increase during acute warming and thus mildly higher Q10-values than seen in controls. Interestingly, the thermally induced limitation of oxygen consumption rate set in earlier in normocapnic than in hypercapnic (1120 µatm CO2) mussels (25.2 °C vs. 28.8 °C), likely due to an onset of metabolic depression in the control group following warming. However, the temperature induced increase in heart rate became limited above 25 °C in both groups indicating an unchanged pejus temperature regardless of CO2 treatment. An upper critical temperature was reached above 28 °C in both treatments indicated by the accumulation of anaerobic metabolites in the mantle tissue, paralleled by a strong increase in haemolymph PCO2 at 31 °C. Ocean acidification caused a decrease in haemolymph pH. The extracellular acidosis remained largely uncompensated despite some bicarbonate accumulation. In all treatments animals developed a progressive warming-induced extracellular acidosis. A stronger pH drop at around 25 °C was followed by stagnating heart rates. However, normocapnic mussels enhanced bicarbonate accumulation at the critical limit, a strategy no longer available to hypercapnic mussels. In conclusion, CO2 has small effects on the response patterns of mussels to warming, leaving thermal thresholds largely unaffected. High resilience of adult North Sea mussels to future ocean acidification indicates that sensitivity to thermal stress is more relevant in shaping the response to future climate change.

Sedimentation and mineral analysis on sediment cores from the Lena Delta

Core and outcrop analysis from Lena mouth deposits have been used to reconstruct the Late Quaternary sedimentation history of the Lena Delta. Sediment properties (heavy mineral composition, grain size characteristics, organic carbon content) and age determinations (14C AMS and IR-OSL) are applied to discriminate the main sedimentary units of the three major geomorphic terraces, which form the delta. The development of the terraces is controlled by complex interactions among the following four factors: (1) Channel migration. According to the distribution of 14C and IR-OSL age determinations of Lena mouth sediments, the major river runoff direction shifted from the west during marine isotope stages 5-3 (third terrace deposits) towards the northwest during marine isotope stage 2 and transition to stage 1 (second terrace), to the northeast and east during the Holocene (first terrace deposits). (2) Eustasy. Sea level rise from Last Glacial lowstand to the modern sea level position, reached at 6-5 ka BP, resulted in back-filling and flooding of the palaeovalleys. (3) Neotectonics. The extension of the Arctic Mid-Ocean Ridge into the Laptev Sea shelf acted as a halfgraben, showing dilatation movements with different subsidence rates. From the continent side, differential neotectonics with uplift and transpression in the Siberian coast ridges are active. Both likely have influenced river behavior by providing sites for preservation, with uplift, in particular, allowing accumulation of deposits in the second terrace in the western sector. The actual delta setting comprises only the eastern sector of the Lena Delta. (4) Peat formation. Polygenetic formation of ice-rich peaty sand (''Ice Complex'') was most extensive (7-11 m in thickness) in the southern part of the delta area between 43 and 14 ka BP (third terrace deposits). In recent times, alluvial peat (5-6 m in thickness) is accumulated on top of the deltaic sequences in the eastern sector (first terrace).

Alkenone-derived UK'37 data and sea surface temperatures (SST) of sediment cores from the Gulf of Taranto (Italy)

Auxiliary data include one file with alkenone-derived UK'37 data and sea surface temperatures (SST). On these data Figs. 7 and 8 of the manuscript are based. The SST are derived from UK'37 by using the transfer function: SST = 29.876 UK'37 - 1.334 of Conte et al. (2006). The data are against the ages (in A.D.) of samples derived from cores GT91-1 (39[deg]59'23"N, 17[deg]45'25"E), GT89-3 and GT90-3 (both 39[deg]45'43"N, 17[deg]53'55"E ). Also included are composite records for UK'37 and SST. For creating the composite records, GT-89-3 was taken as reference core. In the overlapping period the GT89-3 data seem in general lower than the GT91-1 data. To accommodate for this in the composite record, the average difference (0.0343 UK'37 units; equivalent to 1.023 [deg]C) was subtracted from the GT91-1 record. Hereafter, for each depth in the overlapping interval the respective values (UK'37 or SST) of GT89-3 and GT91-1 were averaged. We have also averaged with 16 additional alkenone measurements, from 1793 to 1851, performed in the GT90-3 core.