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.     

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 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.     

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.     

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.     

Appropriate pCO2 treatments in ocean acidification experiments

Experiments in which organisms are reared in treatments simulating current and future pCO₂ concentrations are critical for ocean acidification (OA) research. The majority of OA exposure experiments use average atmospheric pCO₂ levels as a baseline treatment. We conducted an ecoregion-scale analysis of global carbon chemistry datasets. For many locales, atmospheric pCO₂ levels are not an appropriate characterization of marine carbon chemistry. We argue that atmospheric pCO₂ should be disregarded when setting baseline treatment conditions and experimental design should rely on measurements of carbon chemistry in a study subject’s habitat. As carbon chemistry conditions vary with space and time, we suggest using a range of pCO₂ values as a control rather than a single value. We illustrate this issue with data on the habitat of Euphausia pacifica, which currently lives in waters with a pCO₂ around 900 μatm, a concentration much higher than the current global atmospheric mean.     

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.     

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.     

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.     

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.     

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.     

Tough as a rock-boring urchin: adult Echinometra sp. EE from the Red Sea show high resistance to ocean acidification over long-term exposures

Ocean acidification, a process caused by the continuous rise of atmospheric CO₂ levels, is expected to have a profound impact on marine invertebrates. Findings of the numerous studies conducted in this field indicate high variability in species responses to future ocean conditions. This study aimed at understanding the effects of long-term exposure to elevated pCO₂ conditions on the performance of adult Echinometra sp. EE from the Gulf of Aqaba (Red Sea). During an 11-month incubation under high pCO₂ (1,433 μatm, pH_NBS 7.7) and control (435 μatm, pH_NBS 8.1) conditions, we examined the urchins’ somatic and gonadal growth, gametogenesis and skeletal microstructure. Somatic and gonadal growths were exhibited with no significant differences between the treatments. In addition, all urchins in the experiment completed a full reproductive cycle, typical of natural populations, with no detectable impact of increased pCO₂ on the timing, duration or progression of the cycle. Furthermore, scanning electron microscopy imaging of urchin tests and spines revealed no signs of the usual observed effects of acidosis, such as skeletal dissolution, widened stereom pores or non-smoothed structures. Our results, which yielded no significant impact of the high pCO₂ treatment on any of the examined processes in the urchins studied, suggest high resistance of adult Echinometra sp. EE to near future ocean acidification conditions. With respect to other findings in this area, the outcome of this study provides an example of the complicated and diverse responses of echinoids to the predicted environmental changes.     

Ocean acidification induces budding in larval sea urchins

Ocean acidification (OA), the reduction of ocean pH due to hydration of atmospheric CO₂, is known to affect growth and survival of marine invertebrate larvae. Survival and transport of vulnerable planktonic larval stages play important roles in determining population dynamics and community structures in coastal ecosystems. Here, we show that larvae of the purple urchin, Strongylocentrotus purpuratus, underwent high-frequency budding (release of blastula-like particles) when exposed to elevated pCO₂ level (>700 μatm). Budding was observed in >50 % of the population and was synchronized over short periods of time (~24 h), suggesting this phenomenon may be previously overlooked. Although budding can be a mechanism through which larval echinoids asexually reproduce, here, the released buds did not develop into viable clones. OA-induced budding and the associated reduction in larval size suggest new hypotheses regarding physiological and ecological tradeoffs between short-term benefits (e.g. metabolic savings and predation escape) and long-term costs (e.g. tissue loss and delayed development) in the face of climate change.     

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.     

The physiological and molecular responses of larvae from the reef-building coral Pocillopora damicornis exposed to near-future increases in temperature and pCO2

Given the threats of greenhouse gas emissions and a changing climate to marine ecosystems, there is an urgent need to better understand the response of not only adult corals, which are particularly sensitive to environmental changes, but also their larvae, whose mechanisms of acclimation to both temperature increases and ocean acidification are not well understood. Brooded larvae from the reef coral Pocillopora damicornis collected from Nanwan Bay, Southern Taiwan, were exposed to ambient or elevated temperature (25 or 29 °C) and pCO₂ (415 or 635 μatm) in a factorial experiment for 9 days, and a variety of physiological and molecular parameters were measured. Respiration and rubisco protein expression decreased in larvae exposed to elevated temperature, while those incubated at high pCO₂ were larger in size. Collectively, these findings highlight the complex metabolic and molecular responses of this life history stage and the need to integrate our understanding across multiple levels of biological organization. Our results also suggest that for this pocilloporid larval life stage, the impacts of elevated temperature are likely a greater threat under near-future predictions for climate change than ocean acidification.     

The swimming kinematics of larval Atlantic cod, Gadus morhua L., are resilient to elevated seawater pCO2

Kinematics of swimming behavior of larval Atlantic cod, aged 12 and 27 days post-hatch (dph) and cultured under three pCO₂ conditions (control-370, medium-1800, and high-4200 μatm) from March to May 2010, were extracted from swim path recordings obtained using silhouette video photography. The swim paths were analyzed for swim duration, distance and speed, stop duration, and horizontal and vertical turn angles to determine whether elevated seawater pCO₂—at beyond near-future ocean acidification levels—affects the swimming kinematics of Atlantic cod larvae. There were no significant differences in most of the variables tested: the swimming kinematics of Atlantic cod larvae at 12 and 27 dph were highly resilient to extremely elevated pCO₂ levels. Nonetheless, cod larvae cultured at the highest pCO₂ concentration displayed vertical turn angles that were more restricted (median turn angle, 15°) than larvae in the control (19°) and medium (19°) treatments at 12 dph (but not at 27 dph). Significant reduction in the stop duration of cod larvae from the high treatment (median stop duration, 0.28 s) was also observed compared to the larvae from the control group (0.32 s) at 27 dph (but not at 12 dph). The functional and ecological significance of these subtle differences are unclear and, therefore, require further investigation in order to determine whether they are ecologically relevant or spurious.     

Egg and early larval stages of Baltic cod, Gadus morhua, are robust to high levels of ocean acidification

The accumulation of carbon dioxide in the atmosphere will lower the pH in ocean waters, a process termed ocean acidification (OA). Despite its potentially detrimental effects on calcifying organisms, experimental studies on the possible impacts on fish remain scarce. While adults will most likely remain relatively unaffected by changes in seawater pH, early life-history stages are potentially more sensitive, due to the lack of gills with specialized ion-regulatory mechanisms. We tested the effects of OA on growth and development of embryos and larvae of eastern Baltic cod, the commercially most important fish stock in the Baltic Sea. Cod were reared from newly fertilized eggs to early non-feeding larvae in 5 different experiments looking at a range of response variables to OA, as well as the combined effect of CO₂ and temperature. No effect on hatching, survival, development, and otolith size was found at any stage in the development of Baltic cod. Field data show that in the Bornholm Basin, the main spawning site of eastern Baltic cod, in situ levels of pCO₂ are already at levels of 1,100 μatm with a pH of 7.2, mainly due to high eutrophication supporting microbial activity and permanent stratification with little water exchange. Our data show that the eggs and early larval stages of Baltic cod seem to be robust to even high levels of OA (3,200 μatm), indicating an adaptational response to CO₂.     

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.     

Metal contamination increases the sensitivity of larvae but not gametes to ocean acidification in the polychaete Pomatoceros lamarckii (Quatrefages)

Ocean acidification is not happening in isolation but against a background of chronic low-level pollution for most coastal marine environments. The reproductive and larval stages of marine invertebrates can be highly sensitive to the impacts of both environmental pollutants and ocean acidification, but very little is currently known regarding the potential impacts of combined contaminant and high CO₂ exposures on the health of marine organisms. Ocean acidification research to date has focused heavily on the responses of calcifying marine invertebrate larvae and algae, and as such the polychaetes as a group, despite their ecological importance, remain understudied. Here, we investigate the effects of elevated seawater CO₂ (pH range 8.1–7.4, plus an extreme pH of 7.2 in the sperm motility experiments), in combination with the environmental pollutant copper (0.002 μM), on the early life history stages of the intertidal polychaete Pomatoceros lamarckii from two populations. P. lamarckii sperm appear to be robust to elevated seawater CO₂. Whilst all three of the sperm motility end points measured showed a response to elevated CO₂, these responses were small and not linear. The percentage of motile sperm and sperm curvilinear velocity were significantly reduced in the lower pH treatments of 7.4 and 7.2, whereas sperm straight-line velocity (VSL) was mostly unaffected except for an increased VSL at pH 8.0. Fertilisation success was investigated using two populations from the South West (UK), one from Torquay and one from Plymouth Sound. Fertilisation success was slightly but significantly reduced at the 7.6 and 7.4 pH treatments for both populations (a 9.0 % reduction in fertilisation success from pH 8.1 to 7.4 for Torquay), but with a greater effect observed in the population from Plymouth Sound (a 13.33 % reduction in fertilisation success). No additional impact of 0.002 μM copper exposure on fertilisation success was found. Larval survival was found to be much more sensitive to elevated CO₂ than sperm function or fertilisation, and a significant interaction with copper exposure was observed. These results demonstrate the potential for polychaete larvae to be affected by predicted ocean acidification conditions and that chronic coastal pollutants, such as copper, have the potential to alter larval susceptibility to ocean acidification conditions.     

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.