Plio-Pleistocene Mg/Ca and TEX86 sea surface temperature records of ODP Sites 184-1143 and 165-999,

The western warm pools of the Atlantic and Pacific oceans are a critical source of heat and moisture for the tropical climate system. Over the past five million years, global mean temperatures have cooled by 3-4 °C. Yet, current reconstructions of sea surface temperatures indicate that temperature in the warm pools has remained stable during this time. This stability has been used to suggest that tropical sea-surface temperatures are controlled by some sort of thermostat-like regulation. Here we reconstruct sea surface temperatures in the South China Sea, Caribbean Sea and western equatorial Pacific Ocean for the past five million years, using a combination of the Mg/Ca, TEXH86-and Uk'37 surface temperature proxies. Our data indicate that during the period of Pliocene warmth from about 5 to 2.6 million years ago, the western Pacific and western Atlantic warm pools were about 2 °C warmer than today. We suggest that the apparent lack of warming seen in the previous reconstructions was an artefact of low seawater Mg/Ca ratios in the Pliocene oceans. Taking this bias into account, our data indicate that tropical sea surface temperatures did change in conjunction with global mean temperatures. We therefore conclude that the temperature of the warm pools of the equatorial oceans during the Pliocene was not limited by a thermostat-like mechanism.

Temperature dependent growth and inorganic ion composition of haemolymph of juvenile Sepia officinalis, 2006

B Body wet weight and mantle length of juvenile Sepia officinalis were monitored over a peroid of five weeks. The animals had hatched in our aquarium system in Bremerhaven, Germany at 16°C and were descendants of individuals collected in the Oosterschelde estuary, Netherlands. Animals were kept in natural sea water at 10 or 17°C and fed small live shrimp (Palaemonetes varians) ad libitum daily. At the end of the experiment some animals kept at 17°C were sacrificed using ethanol. Haemolymph was withdrawn from the head vein using syringe and needle. Haemolymph samples were stored at -20°C until Na+, Cl-, K+, Mg2+, Ca2+ and SO42- concentrations were determined using ion chromatography. Mean body weight more that tripled at 17°C during the investigation period, while growth was impared by exposue to 10°C. Haemolymph ion concentrations were similar to those in sea water, except for sulphate. The concentration of this ion in the haemolymph was more that ten times lower than in sea water.

Haemolymph inorganic ion composition, physiology and behaviour dependent on temperature and ambient magnesium concentration in early life history stages of Paralomis granulosa (Decapoda, Anomura, Lithodidae)

Marine brachyuran and anomuran crustaceans are completely absent from the extremely cold (-1.8 °C) Antarctic continental shelf, but caridean shrimps are abundant. This has at least partly been attributed to low capacities for magnesium excretion in brachyuran and anomuran lithodid crabs ([Mg2+]HL = 20-50 mmol/L) compared to caridean shrimp species ([Mg2+]HL = 5-12 mmol/L). Magnesium has an anaesthetizing effect and reduces cold tolerance and activity of adult brachyuran crabs. We investigated whether the capacity for magnesium regulation is a factor that influences temperature-dependent activity of early ontogenetic stages of the Sub-Antarctic lithodid crab Paralomis granulosa. Ion composition (Na+, Mg2+, Ca2+, Cl-, [SO4]2-) was measured in haemolymph withdrawn from larval stages, the first and second juvenile instars (crabs I and II) and adult males and females. Magnesium excretion improved during ontogeny, but haemolymph sulphate concentration was lowest in the zoeal stages. Neither haemolymph magnesium concentrations nor Ca2+:Mg2+ ratios paralleled activity levels of the life stages. Long-term (3 week) cold exposure of crab I to 1 °C caused a significant rise of haemolymph sulphate concentration and a decrease in magnesium and calcium concentrations compared to control temperature (9 °C). Spontaneous swimming activity of the zoeal stages was determined at 1, 4 and 9 °C in natural sea water (NSW, [Mg2+] = 51 mmol/L) and in sea water enriched with magnesium (NSW + Mg2+, [Mg2+] = 97 mmol/L). It declined significantly with temperature but only insignificantly with increased magnesium concentration. Spontaneous velocities were low, reflecting the demersal life style of the zoeae. Heart rate, scaphognathite beat rate and forced swimming activity (maxilliped beat rate, zoea I) or antennule beat rate (crab I) were investigated in response to acute temperature change (9, 6, 3, 1, -1 °C) in NSW or NSW + Mg2+. High magnesium concentration reduced heart rates in both stages. The temperature-frequency curve of the maxilliped beat (maximum: 9.6 beats/s at 6.6 °C in NSW) of zoea I was depressed and shifted towards warmer temperatures by 2 °C in NSW + Mg2+, but antennule beat rate of crab I was not affected. Magnesium may therefore influence cold tolerance of highly active larvae, but it remains questionable whether the slow-moving lithodid crabs with demersal larvae would benefit from an enhanced magnesium excretion in nature.

Haemolymph inorganic ion composition, physiology and behaviour dependent on temperature and magnesium in adult Paralomis granulosa (Decapoda, Anomura, Lithodidae)

A low capacity for regulation of extracellular Mg2+ has been proposed to exclude reptant marine decapod crustaceans from temperatures below 0°C and thus to exclude them from the high Antarctic. To test this hypothesis and to elaborate the underlying mechanisms in the most cold-tolerant reptant decapod family of the sub-Antarctic, the Lithodidae, thermal tolerance was determined in the crab Paralomis granulosa (Decapoda, Anomura, Lithodidae) using an acute stepwise temperature protocol (-1°, 1°, 4°, 7°, 10°, and 13°C). Arterial and venous oxygen partial pressures (Po2) in hemolymph, heartbeat and ventilation beat frequencies, and hemolymph cation composition were measured at rest and after a forced activity (righting) trial. Scopes for heartbeat and ventilation beat frequencies and intermittent heartbeat and scaphognathite beat rates at rest were evaluated. Hemolymph [Mg2+] was experimentally reduced from 30 mmol/L to a level naturally observed in Antarctic caridean shrimps (12 mmol/L) to investigate whether the animals remain more active and tolerant to cold (-1°, 1°, and 4°C). In natural seawater, righting speed was significantly slower at -1° and 13°C, compared with acclimation temperature (4°C). Arterial and venous hemolymph Po2 increased in response to cooling even though heartbeat and ventilation beat frequencies as well as scopes decreased. At rest, ionic composition of the hemolymph was not affected by temperature. Activity induced a significant increase in hemolymph [K+] at -1° and 1°C. Reduction of hemolymph [Mg2+] did not result in an increase in activity, an increase in heartbeat and ventilation beat frequencies, or a shift in thermal tolerance to lower temperatures. In conclusion, oxygen delivery in this cold-water crustacean was not acutely limiting cold tolerance, and animals may have been constrained more by their functional capacity and motility. In contrast to earlier findings in temperate and subpolar brachyuran crabs, these constraints remained insensitive to changing Mg2+ levels.

Gill tissue masses and oxygen consumption of Gobionotothen gibberifrons, Notothenia coriiceps and Zoarces viviparus

Mechanisms responsive to hypercapnia (elevated CO2 concentrations) and shaping branchial energy turnover were investigated in isolated perfused gills of two Antarctic Notothenioids (Gobionotothen gibberifrons, Notothenia coriiceps). Branchial oxygen consumption was measured under normo- versus hypercapnic conditions (10,000 ppm CO2) at high extracellular pH values. The fractional costs of ion regulation, protein and RNA synthesis in the energy budgets were determined using specific inhibitors. Overall gill energy turnover was maintained under pH compensated hypercapnia in both Antarctic species as well as in a temperate zoarcid (Zoarces viviparus). However, fractional energy consumption by the examined processes rose drastically in G. gibberifrons (100-180%), and to a lesser extent in N. coriiceps gills (7-56%). In conclusion, high CO2 concentrations under conditions of compensated acidosis induce cost increments in epithelial processes, however, at maintained overall rates of branchial energy turnover.

Temperature tolerance of different larval stages of the spider crab Hyas araneus exposed to elevated seawater PCO2

Exposure to elevated seawater PCO2 limits the thermal tolerance of crustaceans but the underlying mechanisms have not been comprehensively explored. Larval stages of crustaceans are even more sensitive to environmental hypercapnia and possess narrower thermal windows than adults. In a mechanistic approach, we analysed the impact of high seawater CO2 on parameters at different levels of biological organization, from the molecular to the whole animal level. At the whole animal level we measured oxygen consumption, heart rate and activity during acute warming in zoea and megalopa larvae of the spider crab Hyas araneus exposed to different levels of seawater PCO2. Furthermore, the expression of genes responsible for acid-base regulation and mitochondrial energy metabolism, and cellular responses to thermal stress (e.g. the heat shock response) was analysed before and after larvae were heat shocked byrapidly raising the seawater temperature from 10°C rearing temperature to 20°C. Zoea larvae showed a high heat tolerance, which decreased at elevated seawater PCO2, while the already low heat tolerance of megalopa larvae was not limited further by hypercapnic exposure. There was a combined effect of elevated seawater CO2 and heat shock in zoea larvae causing elevated transcript levels of heat shock proteins. In all three larval stages, hypercapnic exposure elicited an up-regulation of genes involved in oxidative phosphorylation, which was, however, not accompanied by increased energetic demands. The combined effect of seawater CO2 and heat shock on the gene expression of heat shock proteins reflects the downward shift in thermal limits seen on the whole animal level and indicates an associated capacity to elicit passive thermal tolerance. The up-regulation of genes involved in oxidative phosphorylation might compensate for enzyme activities being lowered through bicarbonate inhibition and maintain larval standard metabolic rates at high seawater CO2 levels. The present study underlines the necessity to align transcriptomic data with physiological responses when addressing mechanisms affected by an interaction of elevated seawater PCO2 and temperature extremes.

Enzyme activity, body measures and heart mass of Sepia officinalis sampled in the English Channel and Adriatic Sea

In the eurythermal cuttlefish Sepia officinalis, performance depends on hearts that ensure systemic oxygen supply over a broad range of temperatures. We therefore aimed to identify adjustments in energetic cardiac capacity and underlying mitochondrial function supporting thermal acclimation and adaptation that could be crucial for the cuttlefish's competitive success in variable environments. Two genetically distinct cuttlefish populations were acclimated to 11, 16 and 21°C. Subsequently, skinned and permeabilised heart fibres were used to assess mitochondrial functioning by means of high-resolution respirometry and a substrate-inhibitor protocol, followed by measurements of cardiac citrate synthase and cytosolic enzyme activities. Temperate English Channel cuttlefish had lower mitochondrial capacities but larger hearts than subtropical Adriatic cuttlefish. Warm acclimation to 21°C decreased mitochondrial complex I activity in Adriatic cuttlefish and increased complex IV activity in English Channel cuttlefish. However, compensation of mitochondrial capacities did not occur during cold acclimation to 11°C. In systemic hearts, the thermal sensitivity of mitochondrial substrate oxidation was high for proline and pyruvate but low for succinate. Oxygen efficiency of catabolism rose as temperature changed from 11 to 21°C via shifts to oxygen-conserving oxidation of proline and pyruvate and via reduced relative proton leak. The changes observed for substrate oxidation, mitochondrial complexes, relative proton leak and heart mass improve energetic efficiency and essentially seem to extend tolerance to high temperatures and reduce associated tissue hypoxia. We conclude that cuttlefish sustain cardiac performance and, thus, systemic oxygen delivery over short- and long-term changes of temperature and environmental conditions by multiple adjustments in cellular and mitochondrial energetics.

Water temperature data from southern elephant seal from Marion Island with links to datasets

As the Antarctic Circumpolar Current crosses the South-West Indian Ocean Ridge, it creates an extensive eddy field characterised by high sea level anomaly variability. We investigated the diving behaviour of female southern elephant seals from Marion Island during their post-moult migrations in relation to this eddy field in order to determine its role in the animals' at-sea dispersal. Most seals dived within the region significantly more often than predicted by chance, and these dives were generally shallower and shorter than dives outside the eddy field. Mixed effects models estimated reductions of 44.33 ± 3.00 m (maximum depth) and 6.37 ± 0.10 min (dive duration) as a result of diving within the region, along with low between-seal variability (maximum depth: 5.5 % and dive duration: 8.4 %). U-shaped dives increased in frequency inside the eddy field, whereas W-shaped dives with multiple vertical movements decreased. Results suggest that Marion Island's adult female elephant seals' dives are characterised by lowered cost-of-transport when they encounter the eddy field during the start and end of their post-moult migrations. This might result from changes in buoyancy associated with varying body condition upon leaving and returning to the island. Our results do not suggest that the eddy field is a vital foraging ground for Marion Island's southern elephant seals. However, because seals preferentially travel through this area and likely forage opportunistically while minimising transport costs, we hypothesise that climate-mediated changes in the nature or position of this region may alter the seals' at-sea dispersal patterns.

Impact of elevated pCO2 on acid-base regulation of the sea urchin Echinometra mathaei and its relation to resistance to ocean acidification: A study in mesocosms

Due to their low metabolism and apparent poor ion regulation ability, sea urchins could be particularly sensitive to ocean acidification resulting from increased dissolution of atmospheric carbon dioxide. Therefore, we evaluated the acid-base regulation ability of the coral reef sea urchin Echinometra mathaei and the impact of decreased pH on its growth and respiration activity. The study was conducted in two identical artificial reef mesocosms during seven weeks. Experimental tanks were maintained respectively at mean pHT 7.7 and 8.05 (with field-like night and day variations). The major physico-chemical parameters were identical, only pCO2 and pHT differed. Results indicate that E. mathaei can regulate the pH of its coelomic fluid in the considered range of pH, allowing a sustainable growth and ensuring an unaffected respiratory metabolism, at least at short term.

Body sizes and dive characteristics of narwhals (Monodon monoceros) from Kakiak Point and Melville Bay (2005-2007)

We report on wintertime data collected from Baffin Bay and northern Davis Strait, a major gateway linking the Arctic with the subpolar North Atlantic, using narwhals (Monodon monoceros) as an oceanographic sampling platform. Fourteen narwhals were instrumented with satellite-linked time-depth-temperature recorders between 2005 and 2007. Transmitters collected and transmitted water column temperature profiles from each dive between December and April, where >90% of maximum daily dive depths reached the bottom. Temperature measurements were combined with 15 helicopter-based conductivity-temperature-depth (CTD) casts taken in April 2007 across central Baffin Bay and compared with hydrographic climatology values used for the region in Arctic climate models. Winter temperature maxima for whale and CTD data were in good agreement, ranging between 4.0°C and 4.6°C in inshore and offshore Baffin Bay and in Davis Strait. The warm Irminger Water was identified between 57°W and 59°W (at 68°N) between 200 and 400 m depths. Whale data correlated well with climatological temperature maxima; however, they were on average 0.9°C warmer ±0.6°C (P < 0.001). Furthermore, climatology data overestimated the winter surface isothermal layer thickness by 50-80 m. Our results suggest the previously documented warming in Baffin Bay has continued through 2007 and is associated with a warmer West Greenland Current in both of its constituent water masses. This research demonstrates the feasibility of using narwhals as ocean observation platforms in inaccessible Arctic areas where dense sea ice prevents regular oceanographic measurements and where innate site fidelity, affinity for winter pack ice, and multiple daily dives to >1700 m offer a useful opportunity to sample the area.