Ingestion and clearance rates of Copepods

Feeding activity, selective grazing and the potential grazing impact of two dominant grazers of the Polar Frontal Zone, Calanus simillimus and Rhincalanus gigas, and of copepods < 2 mm were investigated with incubation experiments in the course of an iron fertilized diatom bloom in November 2000. All grazers were already actively feeding in the low chlorophyll waters prior to the onset of the bloom. C. simillimus maintained constant clearance rates and fed predominantly on diatoms. R. gigas and the small copepods strongly increased clearance and ingestion of diatoms in response to their enhanced availability. All grazers preyed on microzooplankton, most steadily on ciliates, confirming the view that pure herbivory appears to be the exception rather than the rule in copepod feeding. The grazers exhibited differences in feeding behavior based on selectivity indices. C. simillimus and R. gigas showed prey switching from dinoflagellates to diatoms in response to the phytoplankton bloom. All grazers most efficiently grazed on large diatoms leading to differences in daily losses for large and small species, e.g. Corethron sp. or Thalassionema nitzschioides. Species-specific diatom mortality rates due to grazing suggest that the high feeding activity of C. simillimus prior to and during the bloom played a role in shaping diatom population dynamics

Impact of ocean acidification and warming on the Mediterranean mussel (Mytilus galloprovincialis)

In order to assess the effects of ocean acidification and warming on the Mediterranean mussel (Mytilus galloprovincialis), specimens were reared in aquarium tanks and exposed to elevated conditions of temperature (+3°C) and acidity (-0.3 pH units) for a period of 10 months. The whole system comprised a factorial experimental design with 4 treatments (3 aquaria per treatment): control, lowered pH, elevated temperature, and lowered pH/elevated temperature. Mortality was estimated on a weekly basis and every 2 months, various biometrical parameters and physiological processes were measured: somatic and shell growth, metabolic rates and body fluid acid-base parameters. Mussels were highly sensitive to warming, with 100% mortality observed under elevated temperature at the end of our experiment in October. Mortality rates increased drastically in summer, when water temperature exceeded 25°C. In contrast, our results suggest that survival of this species will not be affected by a pH decrease of 0.3 in the Mediterranean Sea. Somatic and shell growth did not appear very sensitive to ocean acidification and warming during most of the experiment, but were reduced, after summer, in the lowered pH treatment. This was consistent with measured shell net dissolution and observed loss of periostracum, as well as uncompensated extracellular acidosis in the lowered pH treatment indicating a progressive insufficiency in acid-base regulation capacity. However, based on the present dataset, we cannot elucidate if these decreases in growth and regulation capacities after summer are a consequence of lower pH levels during that period or a consequence of a combined effect of acidification and warming. To summarize, while ocean acidification will potentially contribute to lower growth rates, especially in summer when mussels are exposed to sub-optimal conditions, ocean warming will likely pose more serious threats to Mediterranean mussels in this region in the coming decades.

Acid-base regulation in the Dungeness crab (Metacarcinus magister)

Homeostatic regulation allows organisms to secure basic physiological processes in a varying environment. To counteract fluctuations in ambient carbonate system speciation due to elevated seawater pCO2 (hypercapnia), many aquatic crustaceans excrete/accumulate acid-base equivalents through their gills; however, not much is known about the role of ammonia in this response. The present study investigated the effects of hypercapnia on acid-base and ammonia regulation in the Dungeness crab, Metacarcinus magister on the whole animal and isolated gill levels. Hemolymph pCO2 and [HCO3]- increased in M. magister acclimated to elevated pCO2 (330 Pa), while pH remained stable. Additionally, hemolymph [Na+], [Ca2+], and [SO4]2- were significantly increased. When challenged with varying pH during gill perfusion, the pH of the artificial hemolymph remained relatively unchanged. Overall, ammonia production and excretion, as well as oxygen consumption, were reduced in crabs acclimated to elevated pCO2, demonstrating that either (amino acid) oxidation is reduced in response to this particular stress, or nitrogenous wastes are excreted in an alternative form.

Physiological responses and scope for growth upon medium-term exposure to the combined effects of ocean acidification and temperature in a subtidal scavenger Nassarius conoidalis

Physiological responses (ingestion rate, absorption rate and efficiency, respiration, rate, excretion rate) and scope for growth of a subtidal scavenging gastropod Nassarius conoidalis under the combined effects of ocean acidification (pCO2 levels: 380, 950, 1250 µatm) and temperature (15, 30 °C) were investigated for 31 days. There was a significant reduction in all the physiological rates and scope for growth following short-term exposure (1-3 days) to elevated pCO2 except absorption efficiency at 15 °C and 30 °C, and respiration rate and excretion rate at 15 °C. The percentage change in the physiological rates ranged from 0% to 90% at 15 °C and from 0% to 73% at 30 °C when pCO2 was increased from 380 µatm to 1250 µatm. The effect of pCO2 on the physiological rates was enhanced at high temperature for ingestion, absorption, respiration and excretion. When the exposure period was extended to 31 days, the effect of pCO2 was significant on the ingestion rate only. All the physiological rates remained unchanged when temperature increased from 24 °C to 30 °C but the rates at 15 °C were significantly lower, irrespective of the duration of exposure. Our data suggested that a medium-term exposure to ocean acidification has no effect on the energetics of N. conoidalis. Nevertheless, the situation may be complicated by a longer term of exposure and/or a reduction in salinity in a warming world.

Assessing the physiological responses of the gastropod Crepidula fornicata to predicted ocean acidification and warming

Organisms inhabiting coastal waters naturally experience diel and seasonal physico-chemical variations. According to various assumptions, coastal species are either considered to be highly tolerant to environmental changes or, conversely, living at the thresholds of their physiological performance. Therefore, these species are either more resistant or more sensitive, respectively, to ocean acidification and warming. Here, we focused on Crepidula fornicata, an invasive gastropod that colonized bays and estuaries on northwestern European coasts during the 20th century. Small (<3 cm in length) and large (>4.5 cm in length), sexually mature individuals of C. fornicata were raised for 6 months in three different pCO2 conditions (390 µatm, 750 µatm, and 1400 µatm) at four successive temperature levels (10°C, 13°C, 16°C, and 19°C). At each temperature level and in each pCO2 condition, we assessed the physiological rates of respiration, ammonia excretion, filtration and calcification on small and large individuals. Results show that, in general, temperature positively influenced respiration, excretion and filtration rates in both small and large individuals. Conversely, increasing pCO2 negatively affected calcification rates, leading to net dissolution in the most drastic pCO2 condition (1400 µatm) but did not affect the other physiological rates. Overall, our results indicate that C. fornicata can tolerate ocean acidification, particularly in the intermediate pCO2 scenario. Moreover, in this eurythermal species, moderate warming may play a buffering role in the future responses of organisms to ocean acidification.

Impact of medium-term exposure to elevated pCO2 levels on the physiological energetics of the mussel Mytilus chilensis

This study evaluated the impact of medium-term exposure to elevated pCO2 levels (750-1200 ppm) on the physiological processes of juvenile Mytilus chilensis mussels over a period of 70 d in a mesocosm system. Three equilibration tanks filled with filtered seawater were adjusted to three pCO2 levels: 380 (control), 750 and 1200 ppm by bubbling air or an air-CO2 mixture through the water. For the control, atmospheric air (with aprox. 380 ppm CO2) was bubbled into the tank; for the 750 and 1200 ppm treatments, dry air and pure CO2 were blended to each target concentration using mass flow controllers for air and CO2. No impact on feeding activity was observed at the beginning of the experiment, but a significant reduction in clearance rate was observed after 35 d of exposure to highly acidified seawater. Absorption rate and absorption efficiency were reduced at high pCO2 levels. In addition, oxygen uptake fell significantly under these conditions, indicating a metabolic depression. These physiological responses of the mussels resulted in a significant reduction of energy available for growth (scope for growth) with important consequences for the aquaculture of this species during medium-term exposure to acid conditions. The results of this study clearly indicate that high pCO2 levels in the seawater have a negative effect on the health of M. chilensis. Therefore, the predicted acidification of seawater associated with global climate change could be harmful to this ecologically and commercially important mussel.

Seawater acidification affects the physiological energetics and spawning capacity of the Manila clam Ruditapes philippinarum during gonadal maturation

Ocean acidification is predicted to have widespread implications for marine bivalve mollusks. While our understanding of its impact on their physiological and behavioral responses is increasing, little is known about their reproductive responses under future scenarios of anthropogenic climate change. In this study, we examined the physiological energetics of the Manila clam Ruditapes philippinarum exposed to CO2-induced seawater acidification during gonadal maturation. Three recirculating systems filled with 600 L of seawater were manipulated to three pH levels (8.0, 7.7, and 7.4) corresponding to control and projected pH levels for 2100 and 2300. In each system, temperature was gradually increased ca. 0.3 °C per day from 10 to 20 °C for 30 days and maintained at 20 °C for the following 40 days. Irrespective of seawater pH levels, clearance rate (CR), respiration rate (RR), ammonia excretion rate (ER), and scope for growth (SFG) increased after a 30-day stepwise warming protocol. When seawater pH was reduced, CR, ratio of oxygen to nitrogen, and SFG significantly decreased concurrently, whereas ammonia ER increased. RR was virtually unaffected under acidified conditions. Neither temperature nor acidification showed a significant effect on food absorption efficiency. Our findings indicate that energy is allocated away from reproduction under reduced seawater pH, potentially resulting in an impaired or suppressed reproductive function. This interpretation is based on the fact that spawning was induced in only 56% of the clams grown at pH 7.4. Seawater acidification can therefore potentially impair the physiological energetics and spawning capacity of R. philippinarum.

Combined effects of short-term exposure to elevated CO2 and decreased O2 on the physiology and energy budget of the thick shell mussel Mytilus coruscus

Hypoxia and ocean acidification are two consequences of anthropogenic activities. These global trends occur on top of natural variability. In environments such as estuarine areas, short-term acute pH and O2 fluctuations are occurring simultaneously. The present study tested the combined effects of short-term seawater acidification and hypoxia on the physiology and energy budget of the thick shell mussel Mytilus coruscus. Mussels were exposed for 72 h to six combined treatments with three pH levels (8.1, 7.7 and 7.3) and two dissolved oxygen (DO) levels (2 mg/L, 6 mg/L). Clearance rate (CR), food absorption efficiency (AE), respiration rate (RR), ammonium excretion rate (ER), O:N ratio and scope for growth (SFG) were significantly reduced, and faecal organic dry weight ratio (E) was significantly increased at low DO. Low pH did not lead to a reduced SFG. Interactive effects of pH and DO were observed for CR, E and RR. Principal component analysis (PCA) revealed positive relationships among most physiological indicators, especially between SFG and CR under normal DO conditions. These results demonstrate that Mytilus coruscus was sensitive to short-term (72 h) exposure to decreased O2 especially if combined with decreased pH levels. In conclusion, the short-term oxygen and pH variation significantly induced physiological changes of mussels with some interactive effects.

Physiological energetics of the thick shell mussel Mytilus coruscus exposed to seawater acidification and thermal stress

Anthropogenic CO2 emissions have caused seawater temperature elevation and ocean acidification. In view of both phenomena are occurring simultaneously, their combined effects on marine species must be experimentally evaluated. The purpose of this study was to estimate the combined effects of seawater acidification and temperature increase on the energy budget of the thick shell mussel Mytilus coruscus. Juvenile mussels were exposed to six combined treatments with three pH levels (8.1, 7.7 and 7.3) * two temperatures (25 °C and 30 °C) for 14 d. We found that clearance rates (CRs), food absorption efficiencies (AEs), respiration rates (RRs), ammonium excretion rates (ER), scope for growth (SFG) and O:N ratios were significantly reduced by elevated temperature sometimes during the whole experiments. Low pH showed significant negative effects on RR and ER, and significantly increased O:N ratios, but showed almost no effects on CR, AE and SFG of M. coruscus. Nevertheless, their interactive effects were observed in RR, ER and O:N ratios. PCA revealed positive relationships among most physiological indicators, especially between SFG and CR under normal temperatures compared to high temperatures. PCA also showed that the high RR was closely correlated to an increasing ER with increasing pH levels. These results suggest that physiological energetics of juvenile M. coruscus are able to acclimate to CO2 acidification with a little physiological effect, but not increased temperatures. Therefore, the negative effects of a temperature increase could potentially impact the ecophysiological responses of M. coruscus and have significant ecological consequences, mainly in those habitats where this species is dominant in terms of abundance and biomass.

Gut content, fatty acid composition and metabolic rates of the amphipod Anonyx sarsi during POLARSTERN cruise ARK-XIX/1, ice station PS64/070-6

During a winter expedition to the western Barents Sea in March 2003, benthic amphipods of the species Anonyx sarsi were observed directly below pack ice. Only males and juveniles [16.0-37.0 mm long, 16.2-120.8 mg dry mass (DM)] were collected. Guts contained macroalgal fibres, fish eggs and flesh from large carrion. Amphipods had very low levels of total lipids (2.7-17.2% DM). Analysis of lipid biomarkers showed that some of the specimens had preyed on pelagic copepods. Individual respiration rates ranged over 0.4-1.7 ml O2/day (mean: 1.2 ml, SD: 0.5 ml). Individual ammonia excretion rates varied between 7.8 µg and 49.3 µg N/day (mean: 30.7 µg, SD: 15.2 µg). The atomic O:N ratio ranged over 35 to 71 (mean: 55, SD: 14), indicating lipid-dominated metabolism. Mass-specific respiration ranged over 9.8-16.6 ml O2/day/g DM (mean: 13.1 ml, SD: 2.2 ml). The metabolic rates of A. sarsi were twice as high as those of the truly sympagic amphipod Gammarus wilkitzkii, which is better adapted to the under-ice habitat by its energy-saving attached lifestyle. It is concluded that males and juveniles of A. sarsi were actively searching for food in the water column and at the ice underside, but that the nutritional status of the amphipods in late Arctic winter was generally very poor.

Tolerance of juvenile Mytilus galloprovincialis to experimental seawater acidification

Coastal ocean acidification is expected to interfere with the physiology of marine bivalves. In this work, the effects of acidification on the physiology of juvenile mussels Mytilus galloprovincialis were tested by means of controlled CO2 perturbation experiments. The carbonate chemistry of natural (control) seawater was manipulated by injecting CO2 to attain 2 reduced pH levels: -0.3 and -0.6 pH units as compared with the control seawater. After 78 d of exposure, we found that the absorption efficiency and ammonium excretion rate of juveniles were inversely related to pH. Significant differences among treatments were not observed in clearance, ingestion and respiration rates. Coherently, the maximal scope for growth and tissue dry weight were observed in mussels exposed to the pH reduction delta pH=-0.6, suggesting that M. galloprovincialis could be tolerant to CO2 acidification, at least in the highly alkaline coastal waters of Ria Formosa (SW Portugal).