Impact of ocean acidification in the metabolism and swimming behavior of the dolphinfish (Coryphaena hippurus) early larvae

Since the industrial revolution, [CO₂]_atm has increased from 280 μatm to levels now exceeding 380 μatm and is expected to rise to 730–1,020 μatm by the end of this century. The consequent changes in the ocean’s chemistry (e.g., lower pH and availability of the carbonate ions) are expected to pose particular problems for marine organisms, especially in the more vulnerable early life stages. The aim of this study was to investigate how the future predictions of ocean acidification may compromise the metabolism and swimming capabilities of the recently hatched larvae of the tropical dolphinfish (Coryphaena hippurus). Here, we show that the future environmental hypercapnia (ΔpH 0.5; 0.16 % CO₂, ~1,600 μatm) significantly (p < 0.05) reduced oxygen consumption rate up to 17 %. Moreover, the swimming duration and orientation frequency also decreased with increasing pCO₂ (50 and 62.5 %, respectively). We argue that these hypercapnia-driven metabolic and locomotory challenges may potentially influence recruitment, dispersal success, and the population dynamics of this circumtropical oceanic top predator.     

Effects of ocean acidification on the early developmental stages of the horned turban,Turbo cornutus

To estimate the impact of CO₂-driven ocean acidification on the early life stages of gastropods, the effects of increased partial pressure of seawater carbon dioxide (pCO₂) (800–2,000 μatm) on the early developmental stages and larval shell length of the commercially important gastropod, the horned turban snail, Turbo cornutus were investigated. Increase in experimental seawater pCO₂ had an increasingly negative impact on the early developmental rate; the proportion of embryos or larvae displaying retarded development increased at higher pCO₂. The proportion of embryos that developed to the 4-cell stage at 2 h after fertilization decreased linearly with increasing pCO₂. At ~1,000 μatm pCO₂, retarded development was observed in ~50 % of larvae. No embryos developed to the 4-cell stage at 2,000 μatm pCO₂ within 2 h of fertilization. A similar trend continued until 24–26 h after fertilization; the proportion of larvae attaining veliger stage by 24–26 h also decreased with increasing pCO₂. The shell length of T. cornutus veligers decreased gradually as seawater pCO₂ increased, but markedly decreased in seawater under nearly unsaturated and unsaturated conditions (≤1.04) of the aragonite saturation state (Ω _aragonite). The results indicate that increased pCO₂ seawater has a progressive and acute effect on embryonic and larval T. cornutus, and imply that the extended early developmental period and/or the downsized larval shell produced by ocean acidification will have a negative impact on survival, settlement and recruitment well into the future.     

Effects of elevated pCO2 and the effect of parent acclimation on development in the tropical Pacific sea urchin Echinometra mathaei

Effects of acclimation to projected near-future ocean acidification (OA) conditions on physiology, reproduction and development were investigated in the tropical sea urchin Echinometra mathaei. Following 6 weeks in control or one of the three elevated pCO₂ (pH_NIST 7.5–8.1; pCO₂ ~485–1,770 μatm) conditions, adult urchins exhibited a slight decline of growth in low pH treatments and moderately reduced respiration at intermediate levels. At 7 weeks, gametes from adults were used to produce larvae that were reared in their respective parental treatments. To assess whether larvae from acclimated parents are more resilient to elevated pCO₂ than those not acclimated, larvae from control animals were also reared in the elevated pCO₂ treatments. There was no difference in female ‘spawnability’ and oocyte size between treatments, but male spawning ability was reduced in increased pCO₂ conditions. In elevated pCO₂ treatments, the percentage of normal larvae and larval size decreased in the progeny of control- and elevated pCO₂-acclimated parents, and arm asymmetry increased. Thus, acclimation of the parents did not make the progeny more resilient or sensitive to OA effects. Negative effects of increased pCO₂ on reproduction and development may impact on recruitment and population maintenance of this species.     

Effect of increasing sea water pCO2 on the northern Atlantic krill species Nyctiphanes couchii

Surprisingly little is known about potential effects of ocean acidification on krill of the Northern Hemisphere as ecologically very important food web component. Sub-adult individuals of the northern Atlantic krill species Nyctiphanes couchii (caught at Austevoll near Bergen, Norway, in January 2013) were exposed in the laboratory to four different levels of pCO₂ (430, 800, 1,100, and 1,700 µatm) for 5 weeks in order to assess potential changes in a set of biological response variables. Survival decreased and the frequency of moulting-related deaths increased with increasing pCO₂. Survival was considerably reduced at relatively high pCO₂ of 1,700 µatm and tended to be negatively affected at 1,100 µatm pCO₂. However, the experimental results show no significant effects of pCO₂ on inter-moult period and growth at pCO₂ levels below 1,100 µatm. No differences in length measurements of the carapace and uropod were observed across pCO₂ levels, indicating no effect of changing carbonate chemistry on the morphology of those calciferous parts of the exoskeleton. The results suggest that sub-adult N. couchii may not suffer dramatically from predicted near-future changes in pCO₂. However, potential detrimental effects on the moulting process and associated higher mortality at 1,100 µatm pCO₂ cannot be excluded. Further experiments are needed in order to investigate whether early life stages of N. couchii show a different sensitivity to elevated sea water pCO₂ and whether those results are transferable to other krill species of the Northern Hemisphere.     

Photosynthetic and morphological photoacclimation of the seagrass Cymodocea nodosa to season, depth and leaf position

The photoacclimation capacity of the seagrass Cymodocea nodosa was evaluated considering temporal (i.e. seasonal) and spatial (i.e. depth and within-leaf position) factors of variation. Changes along the leaf were measured in a population growing along a depth gradient (from intertidal to subtidal) in Cadiz Bay (Southern Spain) from 2004 to 2005. Photoacclimation was evaluated by photosynthesis (P–E curves), pigment content and leaf morphology. Plants of Cymodocea nodosa showed large physiological and morphological plasticity (mean %CV = 35.8 ± 3.4) according to the three factors considered. Seasonal patterns appeared for photosynthesis, respiration, pigment content and morphology. Nevertheless, seasonal patterns were not consistent with depth or leaf portions. The resulting data set offered different information depending on the analysis conducted; when only one factor (season, depth or leaf portion) was considered, some tendencies observed in the 3-way full design were masked. Accordingly, considering spatio–temporal variability is crucial when describing photoacclimation and estimating productivity in seagrass meadows.     

Ocean acidification and warming alter photosynthesis and calcification of the symbiont-bearing foraminifera Marginopora vertebralis

The impact of elevated CO₂ and temperature on photosynthesis and calcification in the symbiont-bearing benthic foraminifer Marginopora vertebralis was studied. Individual specimens of M. vertebralis were collected from Heron Island on the southern Great Barrier Reef (Australia). They were maintained for 5 weeks at different temperatures (28, 32 °C) and pCO₂ (400, 1,000 µatm) levels spanning a range of current and future climate-change scenarios. The photosynthetic capacity of M. vertebralis was measured with O₂ microsensors and a pulse-amplitude-modulated chlorophyll (Chl) fluorometer, in combination with estimates of Chl a and Chl c ₂ concentrations and calcification rates. After 5 weeks, control specimens remained unaltered for all parameters. Chlorophyll a concentrations significantly decreased in the specimens at 1,000 µatm CO₂ for both temperatures, while no change in Chl c ₂ concentration was observed. Photoinhibition was observed under elevated CO₂ and temperature, with a 70–80 % decrease in the maximum quantum yield of PSII. There was no net O₂ production at elevated temperatures in both CO₂ treatments as compared to the control temperature, supporting that temperature has more impact on photosynthesis and O₂ flux than changes in ambient CO₂. Photosynthetic pigment loss and a decrease in photochemical efficiency are thus likely to occur with increased temperature. The elevated CO₂ and high temperature treatment also lead to a reduction in calcification rate (from +0.1 to >?0.1 % day^?1). Thus, both calcification and photosynthesis of the major sediment-producing foraminifer M. vertebralis appears highly vulnerable to elevated temperature and ocean acidification scenarios predicted in climate-change models.     

Tolerance of juvenile barnacles (Amphibalanus improvisus) to warming and elevated pCO2

We investigated the impacts of warming and elevated pCO₂ on newly settled Amphibalanus improvisus from Kiel Fjord, an estuarine ecosystem characterized by significant natural pCO₂ variability. In two experiments, juvenile barnacles were maintained at two temperature and three pCO₂ levels (20/24 °C, 700–2,140 μatm) for 8 weeks in a batch culture and at four pCO₂ levels (20 °C, 620–2,870 μatm) for 12 weeks in a water flow-through system. Warming as well as elevated pCO₂ hardly affected growth or the condition index of barnacles, although some factor combinations led to temporal significances in enhanced or reduced growth with an increase in pCO₂. While warming increased the shell strength of A. improvisus individuals, elevated pCO₂ had only weak effects. We demonstrate a strong tolerance of juvenile A. improvisus to mean acidification levels of about 1,000 μatm pCO₂ as is already naturally experienced by the investigated barnacle population.     

Influence of sediment nitrogen-availability on carbon and nitrogen dynamics in the seagrass Thalassia testudinum

Variations in tissue carbon (C), nitrogen (N) content, and non-structural carbohydrate (NSC) reserves in the turtle grass Thalassia testudinum Banks ex König were examined in relation to changes in sediment-N availability in Corpus Christi Bay (CCB) and lower Laguna Madre (LLM), Texas, USA, from May to October 1997. Under natural conditions, sediment pore-water NH⁺ ₄-concentrations were higher at CCB (100?μM) than at LLM (30?μM); this difference was reflected in a significantly higher leaf and rhizome N-content at CCB than at LLM. However, sediment NH⁺ ₄-enrichment using a commercial fertilizer resulted in significantly higher tissue N-content relative to controls at both sites. N enrichment also influenced plant carbon metabolism, as reflected by distinct increases in leaf C-content at both sites. Significant decreases in rhizome NSC-content was recorded during the first two months of the experiment, suggesting that C was reallocated from rhizomes to leaves to support stimulated leaf growth at both sites. At LLM, leaf growth-rates increased and leaf turnover-time decreased as a result of sediment NH⁺ ₄-enrichment. With respect to chlorophyll, concentrations did not change significantly at CCB, but increased steadily at LLM after the first month following fertilization. In general however, chlorophyll concentrations in control plots were significantly higher at CCB than that at LLM. These observations suggest that leaf function related to C-fixation is enhanced under higher sediment N conditions, as reflected in higher leaf growth-rates and increased blade chlorophyll-content. In contrast, under low-N conditions, below-ground tissue production is enhanced at the expense of the above-ground shoots and leaves, resulting in the high below: above-ground biomass ratios often observed in seagrass beds of oligotrophic environments.     

Early reproductive stages in the crustose coralline alga Phymatolithon lenormandii are strongly affected by mild ocean acidification

Coralline algae (Corallinales, Rhodophyta) are predicted to be negatively impacted by near-future ocean acidification. The effect of low pH/high pCO₂ on early life stages of Phymatolithon lenormandii (Areschoug) Adey was studied in a perturbation experiment. Several parameters including mortality, calcification (calcein staining) and development (growth and abnormalities) have been monitored for a month under experimental conditions ranging from pH_T = 8.00 (pCO₂ = 398 μatm) and pH_T = 7.55 (pCO₂ = 1,261 μatm). Our results demonstrate that survival and development of P. lenormandii early life stages can be impacted by small pH changes (ΔpH < ?0.1 pH unit). A negative impact of decreasing pH was observed including an increased mortality and a higher rate of abnormalities. Growth and calcification were still observed at the lowest pH (ΔpH = ?0.45). Growth rate was similar at all tested pH, but the maintenance of the skeleton under low pH was only possible through a persistent dynamic dissolution/calcification process, an energetically costly mechanism potentially draining resources from other vital processes.     

Long-term and trans-life-cycle effects of exposure to ocean acidification in the green sea urchin Strongylocentrotus droebachiensis

Anthropogenic CO₂ emissions are acidifying the world’s oceans. A growing body of evidence demonstrates that ocean acidification can impact survival, growth, development and physiology of marine invertebrates. Here, we tested the impact of long-term (up to 16 months) and trans-life-cycle (adult, embryo/larvae and juvenile) exposure to elevated pCO₂ (1,200 μatm, compared to control 400 μatm) on the green sea urchin Strongylocentrotus droebachiensis. Female fecundity was decreased 4.5-fold when acclimated to elevated pCO₂ for 4 months during reproductive conditioning, while no difference was observed in females acclimated for 16 months. Moreover, adult pre-exposure for 4 months to elevated pCO₂ had a direct negative impact on subsequent larval settlement success. Five to nine times fewer offspring reached the juvenile stage in cultures using gametes collected from adults previously acclimated to high pCO₂ for 4 months. However, no difference in larval survival was observed when adults were pre-exposed for 16 months to elevated pCO₂. pCO₂ had no direct negative impact on juvenile survival except when both larvae and juveniles were raised in elevated pCO₂. These negative effects on settlement success and juvenile survival can be attributed to carry-over effects from adults to larvae and from larvae to juveniles. Our results support the contention that adult sea urchins can acclimate to moderately elevated pCO₂ in a matter of a few months and that carry-over effects can exacerbate the negative impact of ocean acidification on larvae and juveniles.     

Future ocean acidification will be amplified by hypoxia in coastal habitats

Ocean acidification is elicited by anthropogenic carbon dioxide emissions and resulting oceanic uptake of excess CO₂ and might constitute an abiotic stressor powerful enough to alter marine ecosystem structures. For surface waters in gas-exchange equilibrium with the atmosphere, models suggest increases in CO₂ partial pressure (pCO₂) from current values of ca. 390 μatm to ca. 700–1,000 μatm by the end of the century. However, in typically unequilibrated coastal hypoxic regions, much higher pCO₂ values can be expected, as heterotrophic degradation of organic material is necessarily related to the production of CO₂ (i.e., dissolved inorganic carbon). Here, we provide data and estimates that, even under current conditions, maximum pCO₂ values of 1,700–3,200 μatm can easily be reached when all oxygen is consumed at salinities between 35 and 20, respectively. Due to the nonlinear nature of the carbonate system, the approximate doubling of seawater pCO₂ in surface waters due to ocean acidification will most strongly affect coastal hypoxic zones as pCO₂ during hypoxia will increase proportionally: we calculate maximum pCO₂ values of ca. 4,500 μatm at a salinity of 20 (T = 10 °C) and ca. 3,400 μatm at a salinity of 35 (T = 10 °C) when all oxygen is consumed. Upwelling processes can bring these CO₂-enriched waters in contact with shallow water ecosystems and may then affect species performance there as well. We conclude that (1) combined stressor experiments (pCO₂ and pO₂) are largely missing at the moment and that (2) coastal ocean acidification experimental designs need to be closely adjusted to carbonate system variability within the specific habitat. In general, the worldwide spread of coastal hypoxic zones also simultaneously is a spread of CO₂-enriched zones. The magnitude of expected changes in pCO₂ in these regions indicates that coastal systems may be more endangered by future global climate change than previously thought.     

Within- and transgenerational effects of ocean acidification on life history of marine three-spined stickleback (Gasterosteus aculeatus)

Some studies have demonstrated that elevated CO₂ concentrations in ocean waters negatively impact metabolism and development of marine fish. Particularly, early developmental stages are probably more susceptible to ocean acidification due to insufficient regulations of their acid-base balance. Transgenerational acclimation can be an important mechanism to mediate impacts of increased CO₂ on marine species, yet very little is known about the potential of parental effects in teleosts. Therefore, transgenerational effects were investigated on life history in juvenile three-spined sticklebacks Gasterosteus aculeatus by acclimating parents (collected in April 2012, 55°03′N, 8°44′E) and offspring to ambient (~400 µatm) and elevated (~1,000 µatm) CO₂ levels and measured parental fecundity as well as offspring survival, growth and otolith characteristics. Exposure to elevated CO₂ concentrations led to an increase in clutch size in adults as well as increased juvenile survival and growth rates between 60 and 90 days post-hatch and enlarged otolith areas compared with fish from ambient CO₂ concentrations. Moreover, transgenerational effects were observed in reduced survival and body size 30 days post-hatch as well as in enlarged otoliths at the end of the experiment, when fathers or both parents were acclimated to the high-CO₂ environment. These results may suggest that elevated CO₂ concentrations had rather positive effects on life-history traits of three-spined sticklebacks, but that parental acclimation can modify these effects without improving offspring fitness. Although the mechanistic basis of such transgenerational acclimation remains unclear, selective gradients within generations seem to determine the direction of transgenerational effects.     

Brooded coral larvae differ in their response to high temperature and elevated pCO2 depending on the day of release

To evaluate the effects of temperature and pCO₂ on coral larvae, brooded larvae of Pocillopora damicornis from Nanwan Bay, Taiwan (21°56.179′N, 120°44.85′E), were exposed to ambient (419–470 μatm) and high (604–742 μatm) pCO₂ at ~25 and ~29 °C in two experiments conducted in March 2010 and March 2012. Larvae were sampled from four consecutive lunar days (LD) synchronized with spawning following the new moon, incubated in treatments for 24 h, and measured for respiration, maximum photochemical efficiency of PSII (F _v/F _m), and mortality. The most striking outcome was a strong effect of time (i.e., LD) on larvae performance: respiration was affected by an LD × temperature interaction in 2010 and 2012, as well as an LD × pCO₂ × temperature interaction in 2012; F _v/F _m was affected by LD in 2010 (but not 2012); and mortality was affected by an LD × pCO₂ interaction in 2010, and an LD × temperature interaction in 2012. There were no main effects of pCO₂ in 2010, but in 2012, high pCO₂ depressed metabolic rate and reduced mortality. Therefore, differences in larval performance depended on day of release and resulted in varying susceptibility to future predicted environmental conditions. These results underscore the importance of considering larval brood variation across days when designing experiments. Subtle differences in experimental outcomes between years suggest that transgenerational plasticity in combination with unique histories of exposure to physical conditions can modulate the response of brooded coral larvae to climate change and ocean acidification.     

Effects of pCO2 on physiology and skeletal mineralogy in a tidal pool coralline alga Corallina elongata

Marine organisms inhabiting environments where pCO₂/pH varies naturally are suggested to be relatively resilient to future ocean acidification. To test this hypothesis, the effect of elevated pCO₂ was investigated in the articulated coralline red alga Corallina elongata from an intertidal rock pool on the north coast of Brittany (France), where pCO₂ naturally varied daily between 70 and 1000 μatm. Metabolism was measured on algae in the laboratory after they had been grown for 3 weeks at pCO₂ concentrations of 380, 550, 750 and 1000 μatm. Net and gross primary production, respiration and calcification rates were assessed by measurements of oxygen and total alkalinity fluxes using incubation chambers in the light and dark. Calcite mol % Mg/Ca (mMg/Ca) was analysed in the tips, branches and basal parts of the fronds, as well as in new skeletal structures produced by the algae in the different pCO₂ treatments. Respiration, gross primary production and calcification in light and dark were not significantly affected by increased pCO₂. Algae grown under elevated pCO₂ (550, 750 and 1000 μatm) formed fewer new structures and produced calcite with a lower mMg/Ca ratio relative to those grown under 380 μatm. This study supports the assumption that C. elongata from a tidal pool, where pCO₂ fluctuates over diel and seasonal cycles, is relatively robust to elevated pCO₂ compared to other recently investigated coralline algae.     

Elevated pCO2 causes developmental delay in early larval Pacific oysters, Crassostrea gigas

Increasing atmospheric CO₂ equilibrates with surface seawater, elevating the concentration of aqueous hydrogen ions. This process, ocean acidification, is a future and contemporary concern for aquatic organisms, causing failures in Pacific oyster (Crassostrea gigas) aquaculture. This experiment determines the effect of elevated pCO₂ on the early development of C. gigas larvae from a wild Pacific Northwest population. Adults were collected from Friday Harbor, Washington, USA (48°31.7′N, 12°1.1′W) and spawned in July 2011. Larvae were exposed to Ambient (400 μatm CO₂), MidCO₂ (700 μatm), or HighCO₂ (1,000 μatm). After 24 h, a greater proportion of larvae in the HighCO₂ treatment were calcified as compared to Ambient. This unexpected observation is attributed to increased metabolic rate coupled with sufficient energy resources. Oyster larvae raised at HighCO₂ showed evidence of a developmental delay by 3 days post-fertilization, which resulted in smaller larvae that were less calcified.     

Environmental impact of CO<Subscript>2</Subscript>, Rn,Hg degassing from the rupture zones produced by Wenchuan <Emphasis Type="Italic">M</Emphasis><Subscript>s</Subscript> 8.0 earthquake in western Sichuan,China

The concentrations and flux of CO₂, ²²²Radon (Rn), and gaseous elemental mercury (Hg) in soil gas were investigated based on the field measurements in June 2010 at ten sites along the seismic rupture zones produced by the May 12, 2008, Wenchuan M _s 8.0 earthquake in order to assess the environmental impact of degassing of CO₂, Rn and Hg. Soil gas concentrations of 344 sampling points were obtained. Seventy measurements of CO₂, Rn and Hg flux by the static accumulation chamber method were performed. The results of risk assessment of CO₂, Rn and Hg concentration in soil gas showed that (1) the concentration of CO₂ in the epicenter of Wenchuan M _s 8.0 earthquake and north end of seismic ruptures had low risk of asphyxia; (2) the concentrations of Rn in the north segment of seismic ruptures had high levels of radon, Maximum was up to level 4, according to Chinese code (GB 50325-2001); (3) the average geoaccumulation index I _geo of soil Hg denoted the lack of soil contamination, and maximum values classified the soil gas as moderately to strongly polluted in the epicenter. The investigation of soil gas CO₂, Rn and Hg degassing rate indicated that (1) the CO₂ in soil gas was characterized by a mean \(\updelta^{13}C_{CO2}\) of ?20.4 ‰ and by a mean CO₂ flux of 88.1 g m^?2 day^?1, which were in the range of the typical values for biologic CO₂ degassing. The maximum of soil CO₂ flux reached values of 399 g m^?2 day^?1 in the epicenter; (2) the soil Rn had higher exhalation in the north segment of seismic ruptures, the maximum reached value of 1976 m Bq m^?2 s^?1; (3) the soil Hg flux was lower, ranging from ?2.5 to 18.7 n g m^?2 h^?1 and increased from south to north. The mean flux over the all profiles was 4.2 n g m^?2 h^?1. The total output of CO₂ and Hg degassing estimated along seismic ruptures for a survey area of 18.17 km² were approximately 0.57 Mt year^?1 and 688.19 g year^?1. It is recommended that land-use planners should incorporate soil gas and/or gas flux measurements in the environmental assessment of areas of possible risk. A survey of all houses along seismic ruptures is advised as structural measures to prevent the ingress of soil gases, including CO₂ and Rn, were needed in some houses.     

Escape performance of temperate king scallop,Pecten maximus under ocean warming and acidification

Among bivalves, scallops are exceptional due to their capacity to escape from predators by swimming which is provided by rapid and strong claps that are produced by the phasic muscle interspersed with tonic muscle contractions. Based on the concept of oxygen and capacity-limited thermal tolerance, the following hypothesis was tested: ocean warming and acidification (OWA) would induce disturbances in aerobic metabolic scope and extracellular acid-case status and impair swimming performance in temperate scallops. Following long-term incubation under near-future OWA scenarios [20 vs. 10 °C (control) and 0.112 kPa CO₂ (hypercapnia) vs. 0.040 kPa CO₂ (normocapnic control)], the clapping performance and metabolic rates (MR) were measured in resting (RMR) and fatigued (maximum MR) king scallops, Pecten maximus, from Roscoff, France. Exposure to OA, either alone or combined with warming, left MR and swimming parameters such as the total number of claps and clapping forces virtually unchanged. Only the duration of the escape response was affected by OA which caused earlier exhaustion in hyper- than in normocapnic scallops at 10 °C. While maximum MR was unaffected, warm exposure increased RMR in both normocapnic and hypercapnic P. maximus resulting in similar Q ₁₀ values of ~2.2. The increased costs of maintenance and the observation of strongly reduced haemolymph PO₂ levels indicate that at 20 °C scallops have reached the upper thermal pejus range with unbalanced capacities for aerobic energy metabolism. As a consequence, warming to 20 °C decreased mean phasic force during escape performance until fatigue. The observed prolonged recovery time in warm incubated scallops might be a consequence of elevated metabolic costs at reduced oxygen availability in the warmth.     

Elevated CO2 affects the behavior of an ecologically and economically important coral reef fish

We tested the effect of near-future CO₂ levels (≈490, 570, 700, and 960 μatm CO₂) on the olfactory responses and activity levels of juvenile coral trout, Plectropomus leopardus, a piscivorous reef fish that is also one of the most important fisheries species on the Great Barrier Reef, Australia. Juvenile coral trout reared for 4 weeks at 570 μatm CO₂ exhibited similar sensory responses and behaviors to juveniles reared at 490 μatm CO₂ (control). In contrast, juveniles reared at 700 and 960 μatm CO₂ exhibited dramatically altered sensory function and behaviors. At these higher CO₂ concentrations, juveniles became attracted to the odor of potential predators, as has been observed in other reef fishes. They were more active, spent less time in shelter, ventured further from shelter, and were bolder than fish reared at 490 or 570 μatm CO₂. These results demonstrate that behavioral impairment of coral trout is unlikely if pCO₂ remains below 600 μatm; however, at higher levels, there are significant impacts on juvenile performance that are likely to affect survival and energy budgets, with consequences for predator–prey interactions and commercial fisheries.     

Effects of nitrogen and phosphorus enrichement on in vivo symbiotic zooxanthellae of Pocillopora damicornis

The reef coral Pocillopora damicornis (Linnaeus) was grown for 8 wk in four nutrient treatments: control, consisting of ambient, unfiltered Kaneohe Bay seawater [dissolved inorganic nitrogen (DIN, 1.0 M) and dissolved inorganic phosphate (DIP, 0.3 M)]; nitrogen enrichment (15 M DIN as ammonium); phosphorus enrichment (1.2 M DIP as inorganic phosphate); and 15 M DIN+1.2 M DIP. Analyses of zooxanthellae for C, N, P and chlorophyll a after the 8 wk experiment indicated that DIN enrichment increased the cellular chlorophyll a and excess nitrogen fraction of the algae, but did not affect C cell^-1. DIP enrichment decreased both C and P cell^-1, but the decrease was proportionally less for C cell^-1. the response of cellular P to both DIN and DIP enrichment appeared to be in the same direction and could not be explained as a primary effect of external nutrient enrichment. The observed response of cellular P might be a consequence of in situ CO₂ limitation. DIN enrichment could increase the CO₂ (aq) demand by increasing the net production per unit area. DIP enrichment could slow down calcification, thus decreasing the availability of CO₂ (aq) in the coral tissue.Hawaii Institute of Marine Biology Contribution No. 920     

The synergistic effects of increasing temperature and CO2 levels on activity capacity and acid–base balance in the spider crab, Hyas araneus

With global climate change, ocean warming and acidification occur concomitantly. In this study, we tested the hypothesis that increasing CO₂ levels affect the acid–base balance and reduce the activity capacity of the Arctic spider crab Hyas araneus, especially at the limits of thermal tolerance. Crabs were acclimated to projected oceanic CO₂ levels for 12 days (today: 380, towards the year 2100: 750 and 1,120 and beyond: 3,000 μatm) and at two temperatures (1 and 4 °C). Effects of these treatments on the righting response (RR) were determined (1) at acclimation temperatures followed by (2) righting when exposed to an additional acute (15 min) heat stress at 12 °C. Prior to (resting) and after the consecutive stresses of combined righting activity and heat exposure, acid–base status and lactate contents were measured in the haemolymph. Under resting conditions, CO₂ caused a decrease in haemolymph pH and an increase in oxygen partial pressure. Despite some buffering via an accumulation of bicarbonate, the extracellular acidosis remained uncompensated at 1 °C, a trend exacerbated when animals were acclimated to 4 °C. The additional combined exposure to activity and heat had only a slight effect on blood gas and acid–base status. Righting activity in all crabs incubated at 1 and 4 °C was unaffected by elevated CO₂ levels or acute heat stress but was significantly reduced when both stressors acted synergistically. This impact was much stronger in the group acclimated at 1 °C where some individuals acclimated to high CO₂ levels stopped responding. Lactate only accumulated in the haemolymph after combined righting and heat stress. In the group acclimated to 1 °C, lactate content was highest under normocapnia and lowest at the highest CO₂ level in line with the finding that RR was largely reduced. In crabs acclimated to 4 °C, the RR was less affected by CO₂ such that activity caused lactate to increase with rising CO₂ levels. In line with the concept of oxygen and capacity limited thermal tolerance, all animals exposed to temperature extremes displayed a reduction in scope for performance, a trend exacerbated by increasing CO₂ levels. Additionally, the differences seen between cold- and warm-acclimated H. araneus after heat stress indicate that a small shift to higher acclimation temperatures also alleviates the response to temperature extremes, indicating a shift in the thermal tolerance window which reduces susceptibility to additional CO₂ exposure.