Posts Tagged 'physiology'

Effects of salinity and pH of seawater on the reproduction of the sea urchin Paracentrotus lividus

Fertilization and early development are usually the most vulnerable stages in the life of marine animals, and the biological processes during this period are highly sensitive to the environment. In nature, sea urchin gametes are shed in seawater, where they undergo external fertilization and embryonic development. In a laboratory, it is possible to follow the exact morphological and biochemical changes taking place in the fertilized eggs and the developing embryos. Thus, observation of successful fertilization and the subsequent embryonic development of sea urchin eggs can be used as a convenient biosensor to assess the quality of the marine environment. In this paper, we have examined how salinity and pH changes affect the normal fertilization process and the following development of Paracentrotus lividus. The results of our studies using confocal microscopy, scanning and transmission electron microscopy, and time-lapse Ca2+ image recording indicated that both dilution and acidification of seawater have subtle but detrimental effects on many aspects of the fertilization process. They include Ca2+ signaling and coordinated actin cytoskeletal changes, leading to a significantly reduced rate of successful fertilization and, eventually, to abnormal or delayed embryonic development.

Continue reading ‘Effects of salinity and pH of seawater on the reproduction of the sea urchin Paracentrotus lividus’

Fluctuating seawater pCO2/pH induces opposing interactions with copper toxicity for two intertidal invertebrates


• Fluctuating pCO2/pH doubled copper-induced antioxidant activity and DNA damage in mussels.

• In contrast, fluctuating conditions mitigated some of the effects of copper for ragworms.

• These effects were associated with the differing acid-base physiology of the two species.

• Physiology was as important as changing copper chemistry in determining overall toxicity.


Global ocean pCO2 is increasing as a result of anthropogenic CO2 emissions, driving a decline in seawater pH. However, coastal waters already undergo fluctuations in pCO2/pH conditions over far shorter timescales, with values regularly exceeding those predicted for the open ocean by the year 2100. The speciation of copper, and therefore its potential toxicity, is affected by changing seawater pH, yet little is known concerning how present-day natural fluctuations in seawater pH affect copper toxicity to marine biota. Here, we test the hypothesis that a fluctuating seawater pCO2/pH regime will alter the responses of the mussel Mytilus edulis and the ragworm Alitta virens to sub-lethal copper, compared to a static seawater pCO2/pH scenario. Mussels and worms were exposed to 0.1 and 0.25 μM copper respectively, concentrations determined to produce comparable toxicity responses in these species, for two weeks under a fluctuating 12-hour pCO2/pH cycle (pH 8.14–7.53, pCO2 445–1747 μatm) or a static pH 8.14 (pCO2 432 μatm) treatment. Mussels underwent a haemolymph acidosis of 0.1–0.2 pH units in the fluctuating treatments, alongside two-fold increases in the superoxide dismutase activity and DNA damage induced by copper, compared to those induced by copper under static pH conditions. Conversely, ragworms experienced an alkalosis of 0.3 pH units under fluctuating pH/pCO2, driven by a two-fold increase in coelomic fluid bicarbonate. This mitigated the copper-induced oxidative stress to slightly reduce both antioxidant activity and DNA damage, relative to the static pH + copper treatment. These opposing responses suggest that differences in species acid-base physiology were more important in determining toxicity responses than the pH-induced speciation change. With variability in seawater chemistry predicted to increase as climate change progresses, understanding how fluctuating conditions interact with the toxicity of pH-sensitive contaminants will become more crucial in predicting their risk to coastal biota.

Continue reading ‘Fluctuating seawater pCO2/pH induces opposing interactions with copper toxicity for two intertidal invertebrates’

Epiphytes provide micro-scale refuge from ocean acidification


• OA induced bleaching and reduced metabolism in non-epiphytized coralline.

• Epiphytized corallines were less susceptible to the detrimental effects of OA.

• Epiphytized corallines had thicker diffusive boundary layer than non-epiphytized.

• Non-calcifying epiphytes provide small scale refuge from OA.

• Epiphytic refugia may protect corallines under future OA conditions.


Coralline algae, a major calcifying component of coastal shallow water communities, have been shown to be one of the more vulnerable taxonomic groups to ocean acidification (OA). Under OA, the interaction between corallines and epiphytes was previously described as both positive and negative. We hypothesized that the photosynthetic activity and the complex structure of non-calcifying epiphytic algae that grow on corallines ameliorate the chemical microenvironmental conditions around them, providing protection from OA. Using mesocosm and microsensor experiments, we showed that the widespread coralline Ellisolandia elongata is less susceptible to the detrimental effects of OA when covered with non-calcifying epiphytic algae, and its diffusive boundary layer is thicker than when not covered by epiphytes. By modifying the microenvironmental carbonate chemistry, epiphytes, facilitated by OA, create micro-scale shield (and refuge) with more basic conditions that may allow the persistence of corallines associated with them during acidified conditions. Such ecological refugia could also assist corallines under near-future anthropogenic OA conditions.

Continue reading ‘Epiphytes provide micro-scale refuge from ocean acidification’

Physiological and ecotoxicological interactions of copper and ocean acidification in the polychaete worms Hediste diversicolor and Alitta virens

For coastal aquatic habitats the change in seawater pH occurring as a result of ocean acidification has the potential to alter the speciation and toxicity of the many contaminants that remain in high concentrations in coastal systems. Of particular concern are metals, such as copper, whose speciation is pH sensitive within the OA range. A meta-analysis of studies to date investigating OA-contaminant interactions using marine invertebrates reveals that 72% of the 44 studies conducted have indeed focused on metals such as copper, with only a few studies looking at polycyclic aromatic hydrocarbons (PAH) and pharmaceuticals. No clear trends in the pH-effect size on contaminant toxicity for either species or contaminant group were present however, suggesting species specific physiological responses may influence this interaction as well as contaminant chemistry. A relatively understudied group were the polychaetes, a key functional group for many coastal sediments. Sediments act as a sink for contaminants where they can accumulate to high concentrations. Hence there is high potential for polychaetes to experience elevated metal exposures under reduced seawater pH as OA progresses. To address this knowledge gap, the responses of two common coastal polychaete, Alitta virens and Hediste diversicolor, were studied under three different experimental scenarios (both water-borne and sediment based) focusing on the physiological and toxicological responses under combined exposures to ocean acidification and copper. Water-borne exposures of Alitta virens to 0.25 μM copper under ambient seawater (pH 8.10) showed a significant increase in DNA damage, along with a rise in both SOD activity and lipid peroxidation. However, when exposed to copper under OA conditions (pH 7.70) there was no further increase in DNA damage and a significant decrease in SOD activity was observed alongside a fall in lipid peroxidation suggesting that OA looks to buffer the toxicity responses to this species. This is in contrast to previous studies using mussels and sea urchins, where copper toxicity responses were significantly higher when exposed under OA conditions. To assess whether local adaptations to high levels of copper contamination influences this OA-copper interaction, a population comparison using a metal resistance population of the harbour ragworm, Hediste diversicolor and a nearby non-resistant population was then conducted. Exposures were run using copper concentrations that elicit comparable toxicity responses, using 0.50 uM copper for the resistant population, compared to 0.25 uM for the non-resistant population, reflecting the two-fold differences in LC50 values for these population. These experiments reveal a significant increase by 19.70% in metabolic rate effect size (the combined stressor when compared to the control) in the resistant population compared to a decrease by 24.02% the non-resistant population, along with differences in ammonia excretion rate and the O:N ratio, thus revealing an energetic cost of this genetic resistance when faced with the combined stressors of OA and copper. These data are in line with the emerging energy limited tolerance to stressors’ hypothesis which states that tolerance to stress can be energy limited, with bioenergetics playing a central role in the tolerance to environmental stress. Finally, a more environmentally realistic exposure scenario was conducted using Alitta virens to test the influence of sediment and tidal cycles on worm acid-base and oxidative stress responses. Field measurements of sediment pH revealed that the pHNBS range over a tidal cycle varies from 6.97 to 7.87, indicating that polychaetes are already experiencing pH’s lower than the predictions for near future open oceans. In aquarium exposures, with overlying water of pHNBS 8.10, sediment pHNBS remained within the range of 7.45 to 7.31, when the overlying water was manipulated to OA conditions (pHNBS 7.70) sediment pHNBS was within the same range as the ambient treatment. The lack of change in sediment pH, despite a 0.40 unit drop in seawater pH, removed any comparative differences in physiological and toxicity end points in the worms between treatments. Tidal emersion induced a slight reduction in sediment pH, with a significant copper effect on sediment pH causing a further decrease in pH levels. Interestingly emersion resulted in a significant OA-copper interaction for coelomic fluid bicarbonate, which increased over the emersion period, however, there was no emersion driven acidosis within coelomic fluid. Overall this work further points to contaminant-OA interactions being species specific driven, in part driven by animal physiology. It also highlights the importance of environmentally relevant exposures with sediment dwelling organisms experiencing lower pH levels than the overlying seawater which could potentially affect metal speciation and could lead to OA-contaminant interactions occurring very differently in this environment. These are important considerations for ecotoxicology studies in the face of global ocean changes.

Continue reading ‘Physiological and ecotoxicological interactions of copper and ocean acidification in the polychaete worms Hediste diversicolor and Alitta virens’

Nutrient availability modulates the effects of climate change on growth and photosynthesis of marine macroalga Pyropia haitanensis (Bangiales, Rhodophyta)

The present research investigated the effect of pCO2 levels (C), seawater temperature (T), and nutrient availability (N) on the growth and physiochemical changes in Pyropia haitanensis. With nutrient enrichment, the interaction of higher pCO2 increased relative growth rates (RGR) by 105.9% when temperature increased (22 °C) compared with the control (lower T, lower C, and lower N: LTLCLN). The higher pCO2 decreased the Pm rates at the lower temperature (18 °C), yet displayed no interaction with higher T or N levels. The higher N increased dark respiration rate (Rd) at 18 °C. At 22 °C, higher pCO2 significantly enhanced the maximum ratio of (quantum yields (Fv/Fo) and the maximum quantum yield (ψpo), while it sharply decreased the absorption of photons per active reaction center (ABS/RC) and dissipation of energy fluxes (per RC) (DIo/RC). Higher temperature obviously reduced the Fv/Fo and ψpo under ambient CO2 level. The higher pCO2 significantly increased the phycoerythrin (PE) and phycocyanin (PC) contents, while higher temperature decreased the PE contents with elevated CO2 and declined the PC content regardless of CO2 condition. At lower nutrient condition, higher pCO2 increased Chl a content. Soluble carbohydrates (SC) and soluble protein (SP) content almost was unchanged among all treatments. Our findings indicate that nutrient availability may regulate photosynthetic mechanism to offset the negative effect of future ocean warming on P. haitanensis, thereby sustaining or increasing the biomass yield of the algae.

Continue reading ‘Nutrient availability modulates the effects of climate change on growth and photosynthesis of marine macroalga Pyropia haitanensis (Bangiales, Rhodophyta)’

Comparative metabolome analysis provides new insights into increased larval mortality under seawater acidification in the sea urchin Strongylocentrotus intermedius


• Seawater acidification’s effects on Strongylocentrotus intermedius larvae were investigated.

• A comparative metabolomics analysis was performed.

• Significantly differentially expressed metabolites were identified and annotated.

• Metabolic pathways related to mortality induced by seawater acidification were observed.


Mortality and metabolic responses of four-armed larvae of Strongylocentrotus intermedius under CO2-induced seawater acidification were investigated. Gametes of S. intermedius were fertilized and developed to the four-armed larval stage in either current natural seawater pH levels (as Control; pH = 7.99 ± 0.01) or laboratory-controlled acidified conditions (OA1: ΔpH = −0.3 units; OA2: ΔpH = −0.4 units; OA3: ΔpH = −0.5 units) according to the predictions of the Intergovernmental Panel on Climate Change (IPCC). The degrees of spicule exposure and asymmetry and mortality of four-armed larvae of S. intermedius were observed; each had a significant linearly increasing trend as the seawater pH level decreased. Comparative metabolome analysis identified a total of 87 significantly differentially expressed metabolites (SDMs, UP: 57, DOWN: 30) in OA-treated groups compared with the control group. Twenty-three SDMs, including carnitine, lysophosphatidylcholine (LPC) 18:3, lysophosphatidyl ethanolamine (LPE) 16:1, glutathione (GSH) and L-ascorbate, exhibited a linear increasing trend with decreasing seawater pH. Nine SDMs exhibited a linear decreasing trend as the seawater pH declined, including hypoxanthine, guanine and thymidine. Among all SDMs, we further mined 48 potential metabolite biomarkers responding to seawater acidification in four-armed larvae of S. intermedius. These potential metabolite biomarkers were mainly enriched in five pathways: glycerophospholipid metabolism, glutathione metabolism, purine metabolism, pyrimidine metabolism and the tricarboxylic acid cycle (TCA cycle). Our results will enrich our knowledge of the molecular mechanisms employed by sea urchins in response to CO2-induced seawater acidification.

Continue reading ‘Comparative metabolome analysis provides new insights into increased larval mortality under seawater acidification in the sea urchin Strongylocentrotus intermedius’

Investigating the effects of climate co-stressors on surf smelt energy demands

Surf smelt (Hypomesus pretiosus) are ecologically and economically important to the Pacific Northwest. They play a critical role in the food web and support numerous commercially important species and are an economically important baitfish. Surf smelt interact closely with the nearshore environment, utilizing approximately 10% of Puget Sound coastlines for spawning throughout the year. Surf smelt spawn at high tide and adhere fertilized eggs to beach sediment, causing their embryos to be exposed to air and seawater throughout embryonic development. Because of this unique life history, surf smelt may be susceptible to anthropogenic stressors including coastal development and climate change. However, very few studies have attempted to test the tolerance of surf smelt to climate change, including elevated temperature and ocean acidification. The purpose of this study was to examine the interactive effects of climate co-stressors ocean acidification and seawater warming on the energy demands of developing surf smelt. Surf smelt embryos and larvae were collected and placed into experimental basins under three temperature treatments (12°C, 15°C, and 18°C) and two total carbon treatments (ambient and elevated) for a period of 14 days for the embryos, and 4 days for the larvae. Increased temperature significantly decreased yolk size in developing surf smelt embryos and larvae. During this time, embryo yolk sacs in the high temperature treatment were on average 10.2% smaller than embryo yolk sacs in ambient temperature water. Larval yolk and oil globules mirrored this trend with larvae in the high temperature treatment having on average 32.5% smaller yolk sacs and 20.0% smaller oil globules compared to larvae in ambient temperature. While no effect of acidification as a singular stressor was observed, the interaction with temperature significantly increased surf smelt embryo heart rates by 5% above ambient conditions. These results indicate that near-future climate change scenarios are going to impact the energy demands of developing surf smelt, a result that may have a variety of potential impacts including altered hatch times, larval deformities, and increased mortality, all of which will increase interannual variability in adult recruitment. The results of this study highlight the need to increase focus on studying surf smelt in the context of ecological and climate change research.

Continue reading ‘Investigating the effects of climate co-stressors on surf smelt energy demands’

Multimarker response of the ragworm Hediste diversicolor (Polychaeta) to seawater acidification derived from potential CO2 leakage from the CCS sub-seabed storage site in the Baltic Sea


• Seawater acidification affected physiological traits, LPO and growth of Hediste diversicolor from the southern Baltic Sea.

• Moderate hypercapnia (pH 7.5–7.1) induced an increase in metabolic rate of the polychaetes and a decline of their growth.

• The most acidic environment (pH 6.5) caused metabolic slow down limiting energy turnover and growth.

• Reduced seawater pH did not impact energetic reserves so, proteins were not used as substrates under acidic conditions.

• High tolerance of the ragworms to hypercapnia stems probably from pre-adaptation to natural pH reduction events in sediment.


Sub-seabed Carbon Capture and Storage (CCS) is conceived as safe technology with small likehood of negative consequences to the marine ecosystem but CO2 escape from geological reservoir still poses potential environmental risk. If carbon dioxide leakage occurs carbonate chemistry in the bottom zone and sessile benthic fauna are expected to be the most likely affected by elevated levels of CO2. Though generic mechanisms and advisory conclusions on the presumable impact of increased acidity on the marine benthic biota were formulated they cannot be applied uniformly across different environmental variables as specific local conditions may alter biological response to hypercapnia. A laboratory experiment was conducted to quantify the effects of medium-term (8 wk) exposure to seawater acidification (pH 7.7–6.5) on the infaunal polychaete Hediste diversicolor from the southern Baltic Sea using multimarker approach. Under moderate acidity (pH 7.5 and 7.1) the polychaetes were found to increase metabolic rate (by 13.4% and 19.6%, respectively) and reduce their body mass (by 8.1% and 5.5% wet weight, respectively and by 6.1% and 3.0% dry weight, respectively) whilst enhancing synthesis of antioxidant malondialdehyde (by 22.8% and 65.3%, respectively). In the most acidic environment (pH 6.5) the ragworms showed overall metabolic slow down (by 34.8%) and impaired growth (e.g. by 10.2% for length of the first three segments) indicative of low vulnerability to hypercapnia. High implicit tolerance of the polychaetes to increased acidity in the environment stems inevitably from a certain level of pre-adaptation to pH reduction events which occur in organic-rich stratified sediments due to intense aerobic biomineralization leading often to oxygen depletion and formation of toxic hydrogen sulphide. Acidification did not affect energetic reserves suggesting that costs of acid-base maintenance were covered mainly from assimilated food and that proteins were not used as metabolic substrates.

Continue reading ‘Multimarker response of the ragworm Hediste diversicolor (Polychaeta) to seawater acidification derived from potential CO2 leakage from the CCS sub-seabed storage site in the Baltic Sea’

Oxidative stress and antioxidant defence responses in two marine copepods in a high CO2 experiment


• Temora revealed higher oxidative stress than Calanus in response to treatment CO2

• Food and predators may have controlled the stress levels, both in fjord and experiment

• Calanus migrating deeper than Temora seems more robust against environmental fluctuations


We collected samples for oxidative stress and antioxidants in a high CO2 mesocosm experiment for two weeks, focussing on two common crustacean copepods Calanus finmarchicus and Temora longicornis. The samples were collected during a field experiment campaign studying responses of plankton communities to future ocean acidification (OA), off the Norwegian coast south of Bergen. The main results showed that there were species-specific differences between Temora and Calanus, especially in antioxidant defences (glutathione system) and oxidative stress (lipid peroxidation and reduced:oxidised glutathione ratio). Regular monitoring of chlorophyll a and jellyfish abundances taking place during the field campaign revealed that both chl a and predators may have affected the eco-physiological response. Antioxidant and oxidative stress levels are known to respond sensitively to both the food quality and quantity and the predator pressure, apart from environmental (i.e., abiotic) changes. Calanus was more robust towards OA, perhaps due to its high tolerance to a wide range of vertical physical-chemical conditions. Both top-down and bottom-up factors seem to play a role for the outcome of copepod responses to future ocean acidification.

Continue reading ‘Oxidative stress and antioxidant defence responses in two marine copepods in a high CO2 experiment’

Physiological feeding rates and cilia suppression in blue mussels (Mytilus edulis) with increased levels of dissolved carbon dioxide


• Increase carbon dioxide decreases cilia beat frequency for blue mussel.

• Ocean acidification decreased clearance rates in blue mussels.

• Ocean acidification resulted in changes in particle selection for marine bivalves.


Gills of marine bivalves, the organs that mediate water flow for feeding and other physiological functions, are exposed to increasing levels of carbon dioxide (CO2) in seawater, in response to ocean acidification (OA). We examined the effects of elevated dissolved CO2 upon filtration and feeding behavior of the blue mussel, Mytilus edulis, under field conditions and in laboratory studies. We further investigated possible changes in cilia beat function in response to elevated dissolved CO2. Physiological filtration and feeding variables measured; included clearance, filtration, organic ingestion, and assimilation rates and selection efficiency, which decreased with increasing CO2. Absorption efficiency was not affected by dissolved CO2. Cilia beat frequency declined in excised lateral cilia (lc) exposed to increasing CO2 levels, which appears to account for decreased clearance rates observed in field and laboratory experiments. Our data suggest that under conditions of increased CO2 blue mussels will experience changes in physiological filtration, feeding rates, and cilia beat function that could have consequences for fitness and performance.

Continue reading ‘Physiological feeding rates and cilia suppression in blue mussels (Mytilus edulis) with increased levels of dissolved carbon dioxide’

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Ocean acidification in the IPCC AR5 WG II

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