Temperature dependence of CO2-enhanced primary production in the European Arctic Ocean

The Arctic Ocean is warming at two to three times the global rate and is perceived to be a bellwether for ocean acidification. Increased CO2 concentrations are expected to have a fertilization effect on marine autotrophs, and higher temperatures should lead to increased rates of planktonic primary production. Yet, simultaneous assessment of warming and increased CO2 on primary production in the Arctic has not been conducted. Here we test the expectation that CO2-enhanced gross primary production (GPP) may be temperature dependent, using data from several oceanographic cruises and experiments from both spring and summer in the European sector of the Arctic Ocean. Results confirm that CO2 enhances GPP (by a factor of up to ten) over a range of 145-2,099?µatm; however, the greatest effects are observed only at lower temperatures and are constrained by nutrient and light availability to the spring period. The temperature dependence of CO2-enhanced primary production has significant implications for metabolic balance in a warmer, CO2-enriched Arctic Ocean in the future. In particular, it indicates that a twofold increase in primary production during the spring is likely in the Arctic.

Seawater carbonate chemistry and physiological response of the Mediterranean crustose coralline alga Lithophyllum cabiochae to elevated pCO2 and temperature

The response of respiration, photosynthesis, and calcification to elevated pCO2 and temperature was investigated in isolation and in combination in the Mediterranean crustose coralline alga Lithophyllum cabiochae. Algae were maintained in aquaria during 1 year at near-ambient conditions of irradiance, at ambient or elevated temperature (+3 °C), and at ambient (ca. 400 µatm) or elevated pCO2 (ca. 700 µatm). Respiration, photosynthesis, and net calcification showed a strong seasonal pattern following the seasonal variations of temperature and irradiance, with higher rates in summer than in winter. Respiration was unaffected by pCO2 but showed a general trend of increase at elevated temperature at all seasons, except in summer under elevated pCO2. Conversely, photosynthesis was strongly affected by pCO2 with a decline under elevated pCO2 in summer, autumn, and winter. In particular, photosynthetic efficiency was reduced under elevated pCO2. Net calcification showed different responses depending on the season. In summer, net calcification increased with rising temperature under ambient pCO2 but decreased with rising temperature under elevated pCO2. Surprisingly, the highest rates in summer were found under elevated pCO2 and ambient temperature. In autumn, winter, and spring, net calcification exhibited a positive or no response at elevated temperature but was unaffected by pCO2. The rate of calcification of L. cabiochae was thus maintained or even enhanced under increased pCO2. However, there is likely a trade-off with other physiological processes. For example, photosynthesis declines in response to increased pCO2 under ambient irradiance. The present study reports only on the physiological response of healthy specimens to ocean warming and acidification, however, these environmental changes may affect the vulnerability of coralline algae to other stresses such as pathogens and necroses that can cause major dissolution, which would have critical consequence for the sustainability of coralligenous habitats and the budgets of carbon and calcium carbonate in coastal Mediterranean ecosystems.

Sea surface temperature oscillations in southern Okinawa Trough during last 2700 years from ODP Hole 195-1202B

Most of the temperature reconstructions for the past two millennia are based on proxy data from various sites on land. Here we present a bidecadal resolution record of sea surface temperature (SST) in Southern Okinawa Trough for the past ca. 2700 years by analyzing tetraether lipids of planktonic archaea in the ODP Hole 1202B, a site under the strong influence of Kuroshio Current and East Asian monsoon. The reconstructed SST anomalies generally coincided with previously reported late Holocene climate events, including the Roman Warm Period, Sui-Tang dynasty Warm Period, Medieval Warm Period, Current Warm Period, Dark Age Cold Period and Little Ice Age. However, the Medieval Warm Period usually thought to be a historical analogue for the Current Warm Period has a mean SST of 0.6-0.8°C lower than that of the Roman Warm Period and Sui-Tang dynasty Warm Period. Despite an increase since 1850 AD, the mean SST in the 20th century is still within the range of natural variability during the past 2700 years. A close correlation of SST in Southern Okinawa Trough with air temperature in East China, intensity of East Asian monsoon and the El-Niño Southern Oscillation index has been attributed to the fluctuations in solar output and oceanic-atmospheric circulation.

Impact of changing carbonate chemistry, temperature, and salinity on growth and test degradation of the benthic foraminifer Ammonia aomoriensis

The present study investigated the combined effects of ocean acidification, temperature, and salinity on growth and test degradation of Ammonia aomoriensis. This species is one of the dominant benthic foraminifera in near-coastal habitats of the southwestern Baltic Sea that can be particularly sensitive to changes in seawater carbonate chemistry. To assess potential responses to ocean acidification and climate change, we performed a fully crossed experiment involving three temperatures (8, 13, and 18°C), three salinities (15, 20, and 25) and four pCO2 levels (566, 1195, 2108, and 3843 µatm) for six weeks. Our results highlight a sensitive response of A. aomoriensis to undersaturated seawater with respect to calcite. The specimens continued to grow and increase their test diameter in treatments with pCO2 <1200 µatm, when Omega calc >1. Growth rates declined when pCO2 exceeded 1200 µatm (Omega calc <1). A significant reduction in test diameter and number of tests due to dissolution was observed below a critical Omega calc of 0.5. Elevated temperature (18°C) led to increased Omega calc, larger test diameter, and lower test degradation. Maximal growth was observed at 18°C. No significant relationship was observed between salinity and test growth. Lowered and undersaturated Omega calc, which results from increasing pCO2 in bottom waters, may cause a significant future decline of the population density of A. aomoriensis in its natural environment. At the same time, this effect might be partially compensated by temperature rise due to global warming.

Experiment: Combined effects of CO2, temperature, irradiance and time on the physiological performance of Chondrus crispus (Rhodophyta)

In natural environments, marine biotas are exposed to a variety of simultaneously acting abiotic factors. Among these, temperature, irradiance and CO2 availability are major factors influencing the physiological performance of marine macroalgae. To test whether elevated levels of CO2 may remediate the otherwise reduced performance of uncalcified seaweeds under the influence of other stressful abiotic factors, we performed multifactorial experiments with the red alga Chondrus crispus from Helgoland (North Sea) with two levels of CO2, temperature and irradiance: low and high pCO2 levels were tested in combination with either (1) optimal and low irradiances or (2) optimal and sub-lethal high temperatures for growth. Performance of C. crispus was evaluated as biomass increase and relative growth rates (RGR), gross photosynthesis and pigment content. Acclimations of growth and photosynthesis were measured after 4 and 8 days. Acclimation time was crucial for elucidating single or combined CO2 effects on growth and photosynthesis. Signifi- cant CO2 effects became evident only in combination with either elevated temperature or reduced irradiance. Growth and photosynthesis had divergent patterns: RGR and biomass significantly increased only under a combination of high pCO2 and elevated temperature; gross photosynthesis was significantly reduced under high pCO2 conditions at low irradiance. Pigment content varied in response to irradiance and temperature, but was independent of pCO2.

Sea-surface temperature reconstruction for sediment core GeoB1023-5

Sediment core GeoB 1023-5 from the eastern South Atlantic was investigated at high temporal resolution for variations of sea-surface temperature (SST) during the past 22 kyr, using the alkenone (UK'37) method. SSTs increased by 3.5°C from about 18°C during the Last Ice Age (21±2 cal kyr BP) to about 21.5°C at 14.5 cal kyr BP. This warming trend associated with the deglaciation phase was followed by a cooling event with lowest SSTs near 20°C, persisting for about 1000 years between 13 and 12 cal kyr BP. The SSTs then continued to increase to about 22.5°C at the Holocene climatic optimum at 7 cal kyr BP, and decreased again during the Late Holocene to a core-top value of 19.8°C that is comparable to modern annual mean SST values. When compared with alkenone SST records from the eastern North Atlantic, our SST record indicates continuous warming throughout the deglaciation phase in the Benguela Current, while its northern counterpart, the Canary Current, experienced prominent cooling during 'Heinrich Event 1' (H1). On the other hand, for the time period corresponding to the 'Younger Dryas' (YD) cooling event, the Benguela SST record exhibits a cold-temperature interval that corresponds to that observed in the eastern North Atlantic SST records. This observation suggests that interhemispheric climate response in Atlantic eastern boundary current systems was different with respect to the two abrupt climate events associated with Termination I. For the H1, the eastern South Atlantic SST record strongly supports the hypothesis that an 'anti-phase' thermal behavior in South Atlantic surface waters was forced by the slowdown of the North Atlantic Deep Water formation during cold spells in the North Atlantic. In contrast, the abrupt cooling in the eastern South Atlantic coincident with the YD period was probably induced by more vigorous global atmospheric circulation, enhancing the upwelling intensity in both eastern boundary current systems. This atmospheric control may have overridden any effect caused by changes in thermohaline circulation on the South Atlantic SSTs during the YD, which leads to the assumption that the thermohaline circulation was already much closer to its interglacial mode during the YD than during the H1.

Sea-surface temperature reconstruction for sediment core GIK18287-3 in the tropical South China Sea

The timing and magnitude of sea-surface temperature (SST) changes in the tropical southern South China Sea (SCS) during the last 16,500 years have been reconstructed on a high-resolution, 14C-dated sediment core using three different foraminiferal transfer functions (SIMMAX28, RAM, FP-12E) and geochemical (Uk'37) SST estimates. In agreement with CLIMAP reconstructions, both the FP-12E and the Uk'37 SST estimates show an average late glacial-interglacial SST difference of 2.0°C, whereas the RAM and SIMMAX28 foraminiferal transfer functions show only a minor (0.6°C) or no consistent late glacial-interglacial SST change, respectively. Both the Uk'37 and the FP-12E SST estimates, as well as the planktonic foraminiferal delta18O values, indicate an abrupt warming (ca. 1°C in <200 yr) at the end of the last glaciation, synchronous (within dating uncertainties) with the Bølling transition as recorded in the Greenland Ice Sheet Project 2 (GISP2) ice core, whereas the RAM-derived deglacial SST increase appears to lag during this event by ca. 500 yr. The similarity in abruptness and timing of the warming associated with the Bølling transition in Greenland and the southern SCS suggest a true synchrony of the Northern Hemisphere warming at the end of the last glaciation. In contrast to the foraminiferal transfer function estimates that do not indicate any consistent cooling associated with the Younger Dryas (YD) climate event in the tropical SCS, the Uk'37 SST estimates show a cooling of ca. 0.2-0.6°C compared to the Bølling-Allerød period. These Uk'37 SST estimates from the southern SCS argue in favor of a Northern Hemisphere-wide, synchronous cooling during the YD period.

Sea surface temperature reconstruction for the Southwest Pacific Ocean

Paleoceanographic archives derived from 17 marine sediment cores reconstruct the response of the Southwest Pacific Ocean to the peak interglacial, Marine Isotope Stage (MIS) 5e (ca. 125?ka). Paleo-Sea Surface Temperature (SST) estimates were obtained from the Random Forest model-an ensemble decision tree tool-applied to core-top planktonic foraminiferal faunas calibrated to modern SSTs. The reconstructed geographic pattern of the SST anomaly (maximum SST between 120 and 132?ka minus mean modern SST) seems to indicate how MIS 5e conditions were generally warmer in the Southwest Pacific, especially in the western Tasman Sea where a strengthened East Australian Current (EAC) likely extended subtropical influence to ca. 45°S off Tasmania. In contrast, the eastern Tasman Sea may have had a modest cooling except around 45°S. The observed pattern resembles that developing under the present warming trend in the region. An increase in wind stress curl over the modern South Pacific is hypothesized to have spun-up the South Pacific Subtropical Gyre, with concurrent increase in subtropical flow in the western boundary currents that include the EAC. However, warmer temperatures along the Subtropical Front and Campbell Plateau to the south suggest that the relative influence of the boundary inflows to eastern New Zealand may have differed in MIS 5e, and these currents may have followed different paths compared to today.

Surface temperature reconstruction using alkenones from sediment core B0406 retreived in the Peru upwelling area

We reconstructed a high-resolution, alkenone-based sea surface temperature (SST) record spanning the last ca. 150 years, from a sediment core retrieved within the main upwelling zone off Peru. A conspicuous SST decline is evidenced since the 1950s despite interdecadal SST variability. Instrumental SST data and reanalysis of ECMWF ERA 40 winds suggest that the recent coastal cooling corresponds mainly to an intensification of alongshore winds and associated increase of upwelling in spring. Consistently, both proxy and instrumental data evidence increased productivity in phase with the SST cooling. Our data expand on previous reports on recent SST cooling in other Eastern Boundary upwelling systems and support scenarios that relate coastal upwelling intensification to global warming. Yet, further investigations are needed to assess the role of different mechanisms and forcings (enhanced local winds vs. spin-up of the South Pacific High Pressure cell).

Experiment: Elevated temperature and PCO2 affect enzyme activities in differentially oxidative tissues of Notothenia rossii

Mitochondrial plasticity plays a central role in setting the capacity for acclimation of aerobic metabolism in ectotherms in response to environmental changes. We still lack a clear picture if and to what extent the energy metabolism and mitochondrial enzymes of Antarctic fish can compensate for changing temperatures or PCO2 and whether capacities for compensation differ between tissues. We therefore measured activities of key mitochondrial enzymes (citrate synthase (CS), cytochrome c oxidase (COX)) from heart, red muscle, white muscle and liver in the Antarctic fish Notothenia rossii after warm- (7 °C) and hypercapnia- (0.2 kPa CO2) acclimation vs. control conditions (1 °C, 0.04 kPa CO2). In heart, enzymes showed elevated activities after cold-hypercapnia acclimation, and a warm-acclimation-induced upward shift in thermal optima. The strongest increase in enzyme activities in response to hypercapnia occurred in red muscle. In white muscle, enzyme activities were temperature-compensated. CS activity in liver decreased after warm-normocapnia acclimation (temperature-compensation), while COX activities were lower after cold- and warm-hypercapnia exposure, but increased after warm-normocapnia acclimation. In conclusion, warm-acclimated N. rossii display low thermal compensation in response to rising energy demand in highly aerobic tissues, such as heart and red muscle. Chronic environmental hypercapnia elicits increased enzyme activities in these tissues, possibly to compensate for an elevated energy demand for acid-base regulation or a compromised mitochondrial metabolism, that is predicted to occur in response to hypercapnia exposure. This might be supported by enhanced metabolisation of liver energy stores. These patterns reflect a limited capacity of N. rossii to reorganise energy metabolism in response to rising temperature and PCO2.

(Table 1) TOC content, BIT, TEX86 and UK'37 indices, and sea surface temperature reconstruction for the middle Miocene to Pliocene of ODP Hole 175-1085A

The initiation of the Benguela upwelling has been dated to the late Miocene, but estimates of its sea surface temperature evolution are not available. This study presents data from Ocean Drilling Program (ODP) Site 1085 recovered from the southern Cape Basin. Samples of the middle Miocene to Pliocene were analyzed for alkenone-based (UK'37, SSTUK) and glycerol dialkyl glycerol tetraether (GDGT) based (TEX86, TempTEX) water temperature proxies. In concordance with global cooling during the Miocene, SSTUK and TempTEX exhibit a decline of about 8°C and 16°C, respectively. The temperature trends suggest an inflow of cold Antarctic waters triggered by Antarctic ice sheet expansion and intensification of Southern Hemisphere southeasterly winds. A temperature offset between both proxies developed with the onset of upwelling, which can be explained by differences in habitat: alkenone-producing phytoplankton live in the euphotic zone and record sea surface temperatures, while GDGT-producing Thaumarchaeota are displaced to colder subsurface waters in upwelling-influenced areas and record subsurface water temperatures. We suggest that variations in subsurface water temperatures were driven by advection of cold Antarctic waters and thermocline adjustments that were due to changes in North Atlantic deep water formation. A decline in surface temperatures, an increased offset between temperature proxies, and an increase in primary productivity suggest the establishment of the Benguela upwelling at 10 Ma. During the Messinian Salinity Crisis, between 7 and 5 Ma, surface and subsurface temperature estimates became similar, likely because of a strong reduction in Atlantic overturning circulation, while high total organic carbon contents suggest a "biogenic bloom." In the Pliocene the offset between the temperature estimates and the cooling trend was reestablished.

Sea surface temperature estimation of sediment core 86014-12PC51

A recently developed technique for determining past sea surface temperatures (SST), based on an analysis of the unsaturation ratio of long chain C37 methyl alkenones produced by Prymnesiophyceae phytoplankton (U37 k' ), has been applied to an upper Quaternary sediment core from the equatorial Atlantic. U37 k' temperature estimates were compared to those obtained from delta18O of the planktonic foraminifer Globigerinoides sacculifer and of planktonic foraminiferal assemblages for the last glacial cycle. The alkenone method showed 1.8°C cooling at the last glacial maximum, about 1/2 to 1/3 of the decrease shown by the isotopic method (6.3°C) and foraminiferal modern analogue technique estimates for the warm season (3.8°C). Warm season foraminiferal assemblage estimates based on transfer functions are out of phase with the other estimates, showing a 1.4°C drop at the last glacial maximum with an additional 0.9°C drop in the deglaciation. Increased alkenone abundances, total organic carbon percentage and foraminiferal accumulation rates in the last glaciation indicate an increase in productivity of as much as 4 times over present day. These changes are thought to be due to increased upwelling caused by enhanced winds during the glaciation. If U37 k' estimates are correct, as much as 50-70% (up to 4.5°C) of estimated delta18O and modern analogue temperature changes in the last glaciation may have been due to changes in thermocline depth, whereas transfer functions seem more strongly influenced by seasonality changes. This indicates these estimates may be influenced as strongly by other factors as they are by SST, which in the equatorial Atlantic was only reduced slightly in the last glaciation.

The larvae of congeneric gastropods showed differential responses to the combined effects of ocean acidification, temperature and salinity

The tolerance and physiological responses of the larvae of two congeneric gastropods, the intertidal Nassarius festivus and subtidal Nassarius conoidalis, to the combined effects of ocean acidification (PCO2 at 380, 950, 1250 ppm), temperature (15, 30 degrees C) and salinity (10, 30 psu) were compared. Results of three-way ANOVA on cumulative mortality after 72-h exposure showed significant interactive effects in which mortality increased with pCO(2) and temperature, but reduced at higher salinity for both species, with higher mortality being obtained for N. conoidalis. Similarly, respiration rate of the larvae increased with temperature and pCO(2) level for both species, with a larger percentage increase for N. conoidalis. Larval swimming speed increased with temperature and salinity for both species whereas higher pCO(2) reduced swimming speed in N. conoidalis but not N. festivus. The present findings indicated that subtidal congeneric species are more sensitive than their intertidal counterparts to the combined effects of these stressors. (c) 2014 Elsevier Ltd. All rights reserved.

(Table S2) The reconstructed sea surface temperature (SST) over the eastern equatorial Pacific

Abundant hydroclimatic evidence from western Amazonia and the adjacent Andes documents wet conditions during Heinrich Stadial 1 (HS1, 18-15 ka), a cold period in the high latitudes of the North Atlantic. This precipitation anomaly was attributed to a strengthening of the South American summer monsoon due to a change in the Atlantic interhemispheric sea surface temperature (SST) gradient. However, the physical viability of this mechanism has never been rigorously tested. We address this issue by combining a thorough compilation of tropical South American paleorecords and a set of atmosphere model sensitivity experiments. Our results show that the Atlantic SST variations alone, although leading to dry conditions in northern South America and wet conditions in northeastern Brazil, cannot produce increased precipitation over western Amazonia and the adjacent Andes during HS1. Instead, an eastern equatorial Pacific SST increase (i.e., 0.5-1.5 °C), in response to the slowdown of the Atlantic Meridional Overturning Circulation during HS1, is crucial to generate the wet conditions in these regions. The mechanism works via anomalous low sea level pressure over the eastern equatorial Pacific, which promotes a regional easterly low-level wind anomaly and moisture recycling from central Amazonia towards the Andes.