The continuing increase of carbon dioxide (CO2) levels in the atmosphere leads to increases in global temperatures and partial pressure of CO2 (PCO2) in surface waters, causing ocean acidification. These changes are especially pronounced in shallow coastal and estuarine waters and are expected to significantly affect marine calcifiers including bivalves that are ecosystem engineers in estuarine and coastal communities. To elucidate potential effects of higher temperatures and PCO2 on physiology and biomineralization of marine bivalves, we exposed two bivalve species, the eastern oysters Crassostrea virginica and the hard clams Mercenaria mercenaria to different combinations of PCO2 (~ 400 and 800 μatm) and temperatures (22 and 27 °C) for 15 weeks. Survival, bioenergetic traits (tissue levels of lipids, glycogen, glucose and high energy phosphates) and biomineralization parameters (mechanical properties of the shells and activity of carbonic anhydrase, CA) were determined in clams and oysters under different temperature and PCO2 regimes. Our analysis showed major inter-species differences in shell mechanical traits and bioenergetics parameters. Elevated temperature led to the depletion of tissue energy reserves indicating energy deficiency in both species and resulted in higher mortality in oysters. Interestingly, while elevated PCO2 had a small effect on the physiology and metabolism of both species, it improved survival in oysters. At the same time, a combination of high temperature and elevated PCO2 lead to a significant decrease in shell hardness in both species, suggesting major changes in their biomineralization processes. Overall, these studies show that global climate change and ocean acidification might have complex interactive effects on physiology, metabolism and biomineralization in coastal and estuarine marine bivalves.
Posts Tagged 'laboratory'
Interactive effects of elevated temperature and CO2 levels on energy metabolism and biomineralization of marine bivalves Crassostrea virginica and Mercenaria mercenariaPublished 24 May 2013 Science Leave a Comment
Tags: biological response, laboratory, mollusks, morphology, mortality, multiple factors, physiology, temperature
Physiological compensation for environmental acidification is limited in the deep-sea urchin Strongylocentrotus fragilisPublished 22 May 2013 Science Leave a Comment
Tags: biological response, echinoderms, laboratory, morphology, mortality, multiple factors, North Pacific, oxygen, performance, physiology
Anthropogenic CO2 is now reaching depths over 1000 m in the Eastern Pacific, overlapping the Oxygen Minimum Zone (OMZ). Deep-sea animals – particularly, calcifiers – are suspected to be especially sensitive to environmental acidification associated with global climate change. We have investigated the effects of hypercapnia and hypoxia on the deep-sea urchin Strongylocentrotus fragilis, during two long-term exposure experiments (1 month and 4 month) at three levels of reduced pH at in situ O2 levels of approx. 10% saturation, and also to control pH at 100% O2 saturation. During the first experiment, internal acid-base balance was investigated during a one-month exposure; results show S. fragilis has limited ability to compensate for the respiratory acidosis brought on by reduced pH, due in part to low non-bicarbonate extracellular fluid buffering capacity. During the second experiment, longer-term effects of hypercapnia and variable O2 on locomotion, feeding, growth, and gonadosomatic index (GSI) were investigated; results show significant mortality and correlation of all measured parameters with environmental acidification at pH 6.6. Transient adverse effects on locomotion and feeding were seen at pH 7.2, without compromise of growth or GSI. Based on the expected changes in ocean pH and oxygen, results suggest extinction of S. fragilis in the eastern North Pacific is unlikely. Rather, we expect a shoaling and contraction of its bathymetric range.
CO2-driven ocean acidification reduces larval feeding efficiency and change food selectivity in the mollusk Concholepas concholepasPublished 22 May 2013 Science Leave a Comment
Tags: biological response, laboratory, mollusks, performance, South Pacific
We present experimental data obtained from an experiment with newly hatched veliger larvae of the gastropod Concholepas concholepas exposed to three pCO2 levels. Egg capsules were collected from two locations in northern and central Chile, and then incubated throughout their entire intra-capsular life cycle at three nominal pCO2 levels, ∼400, 700 and 1000 ppm (i.e. corresponding to ∼8.0, 7.8 and 7.6 pH units, respectively). Hatched larvae were fed with natural food assemblages. Food availability at time zero did not vary significantly with pCO2 level. Our results clearly showed a significant effect of elevated pCO2 on the intensity of larval feeding, which dropped by >60%. Incubation also showed that pCO2-driven ocean acidification (OA) may radically impact the selectivity of ingested food by C. concholepas larvae. Results also showed that larvae switched their clearance rate based on large cells, such as diatoms and dinoflagellates to tiny and highly abundant nanoflagellates and cyanobacteria as pCO2 levels increased. Thus, this study reveals the important effect of low pH conditions on larval feeding behavior, in terms of both ingestion magnitude and selectivity. These findings support the notion that larval feeding is a key physiological process susceptible to the effects of OA.
Tags: biological response, crustaceans, laboratory, multiple factors, physiology, temperature
The injection of anthropogenically-produced CO2 into the atmosphere will lead to an increase in temperature and a decrease in pH at the surface of the oceans by 2100. Marine intertidal organisms possess the ability to cope in the short term with environmental fluctuations exceeding predicted values. However, how they will cope with chronic exposure to elevated temperature and pCO2 is virtually unknown. In addition, individuals from the same species/population often show remarkable levels of variation in their responses to complex climatic changes: in particular, variation in metabolic rates often is linked to differences in individuals’ performances and fitness. Despite its ecological and evolutionary importance, inter-individual variation has rarely been investigated within the context of climatic changes, and most investigations have typically employed orthogonal experimental designs paired to analyses of independent samples. Although this is undoubtedly a powerful and useful approach, it may not be the most appropriate for understanding all alterations of biological functions in response to environmental changes. An individual approach arguably should be favored when trying to describe organisms’ responses to climatic change. Consequently, to test which approach had the greater power to discriminate the intensity and direction of an organism’s response to complex climatic changes, we investigated the extracellular osmo/iono-regulatory abilities, upper thermal tolerances (UTTs), and metabolic rates of individual adults of an intertidal amphipod, Echinogammarus marinus, exposed for 15 days to combined elevated temperature and pCO2. The individual approach led to stronger and different predictions on how ectotherms will likely respond to ongoing complex climatic change, compared with the independent approaches. Consequently, this may call into question the relevance, or even the validity, of some of the predictions made to date. Finally, we argue that treating individual differences as biologically meaningful can lead to a better understanding of the physiological responses themselves and the selective processes that will occur with complex climatic changes; selection will likely play a crucial role in defining species’ responses to future environmental changes. Individuals with higher metabolic rates were also characterized by greater extracellular osmo/iono-regulative abilities and higher UTTs, and thus there appeared to be no evolutionary trade-offs between these functions. However, as individuals with greater metabolic rates also have greater costs for maintenance and repair, and likely a lower fraction of energy available for growth and reproduction, trade-offs between life-history and physiological performance may still arise.
Effects of ocean warming and acidification on survival, growth and skeletal development in the early benthic juvenile sea urchin (Heliocidaris erythrogramma)Published 22 May 2013 Science Leave a Comment
Tags: biological response, echinoderms, laboratory, morphology, multiple factors, survival, temperature
Co-occurring ocean warming, acidification and reduced carbonate mineral saturation have significant impacts on marine biota, especially calcifying organisms. The effects of these stressors on development and calcification in newly metamorphosed juveniles (ca. 0.5 mm test diam) of the intertidal sea urchin Heliocidaris erythrogramma, an ecologically important species in temperate Australia, were investigated in context with present and projected future conditions. Habitat temperature and pH/pCO2 were documented to place experiments in a biologically and ecologically relevant context. These parameters fluctuated diurnally up to 10°C and 0.45 pH units. The juveniles were exposed to three temperature (21°C, 23°C, 25°C) and four pH (8.1, 7.8, 7.6, 7.4) treatments in all combinations, representing ambient sea surface conditions (21°C, pH 8.1; pCO2 397; ΩCa 4.7; ΩAr 3.1), near-future projected change (+2-4°C, -0.3-0.5 pH units; pCO2 400-1820; ΩCa 5.0-1.6; ΩAr 3.3-1.1), and extreme conditions experienced at low tide (+4°C, -0.3-07 pH units; pCO2 2850-2967; ΩCa 1.1-1.0; ΩAr 0.7-0.6). The lowest pH treatment (pH 7.4) was used to assess tolerance levels. Juvenile survival and test growth were resilient to current and near-future warming and acidification. Spine development, however, was negatively effected by near-future increased temperature (+2-4°C) and extreme acidification (pH 7.4), with a complex interaction between stressors. Near-future warming was the more significant stressor. Spine tips were dissolved in the pH 7.4 treatments. Adaptation to fluctuating temperature-pH conditions in the intertidal may convey resilience to juvenile H. erythrogramma to changing ocean conditions, however, ocean warming and acidification may shift baseline intertidal temperature and pH/pCO2 to levels that exceed tolerance limits.
Results of laboratory and field experiments of the direct effect of increasing CO2 on net primary production of macroalgal species in brackish-water ecosystemsPublished 22 May 2013 Science Leave a Comment
Tags: algae, Baltic Sea, biological response, field, laboratory, mesocosms, primary production
Studies on the effects of increasing acidification on marine communities have been previously mostly carried out in truly marine areas whereas brackish-water ecosystems such as the Baltic Sea have been less studied. The current study analyses how acidification induced by elevated atmospheric carbon dioxide affects the photosynthetic net production of different macroalgal species in the brackish Baltic Sea. Research methods include sets of laboratory and field experiments carried out in shallow coastal brackish waters. The aim of the laboratory experiments was to develop the necessary techniques and experience for the mesocosm experiments. Laboratory experiments were carried out using specimens of the red alga Furcellaria lumbricalis collected from Kakumäe Bay. The mesocosm experiments were conducted in Kõiguste Bay during the field season of 2011. Separate mesocosms were operated in each set with different CO2 concentrations and a control treatment in natural conditions. Field experiments were carried out with three species representing three different morphological and ecological groups: Ulva intestinalis, a fast-growing green alga; Fucus vesiculosus, a perennial brown alga with a slow metabolism; and Furcellaria lumbricalis, a perennial red alga. Photosynthetic activity was used as the response variable. In the laboratory decreasing pH increased the net primary production of F. lumbricalis with the lowest net primary production values measured at pH 8.0 and the highest at pH 6.5. Results of the field experiments indicated that increased CO2 levels in seawater favoured photosynthetic activity of the macroalgae U. intestinalis and F. lumbricalis, but F. vesiculosus showed no response to elevated CO2. Elevated CO2 levels are suggested to favour the production of fast-growing filamentous species, which thus may indirectly enhance the effect of eutrophication in the shallow coastal brackish waters.
Natural variation, and the capacity to adapt to ocean acidification in the keystone sea urchin Strongylocentrotus purpuratusPublished 17 May 2013 Science Leave a Comment
Tags: adaptation, biological response, echinoderms, laboratory, molecular biology, morphology, North Pacific
A rapidly growing body of literature documents the potential negative effects of CO2-driven ocean acidification (OA) on marine organisms. However, nearly all of this work has focused on the effects of future conditions on modern populations, neglecting the role of adaptation. Rapid evolution can alter demographic responses to environmental change, ultimately affecting the likelihood of population persistence, but the capacity for adaptation will differ among populations and species. Here, we measure the capacity of the ecologically important purple sea urchin Strongylocentrotus purpuratus to adapt to OA, using a breeding experiment to estimate additive genetic variance for larval size (an important component of fitness) under future high pCO2/low pH conditions. Although larvae reared under future conditions were smaller than those reared under present-day conditions, we show that there is also abundant genetic variation for body size under elevated pCO2, indicating that this trait can evolve. The observed heritability of size was 0.40±0.32 (95% CI) under low pCO2, and 0.50±0.30 under high pCO2 conditions. Accounting for the observed genetic variation in models of future larval size and demographic rates substantially alters projections of performance for this species in the future ocean. Importantly, our model shows that after incorporating the effects of adaptation, the OA-driven decrease in population growth rate is up to 50% smaller, than that predicted by the “no-adaptation” scenario. Adults used in the experiment were collected from two sites on the coast of the Northeast Pacific that are characterized by different pH regimes, as measured by autonomous sensors. Comparing results between sites, we also found subtle differences in larval size under high pCO2 rearing conditions, consistent with local adaptation to carbonate chemistry in the field. These results suggest that spatially varying selection may help to maintain genetic variation necessary for adaptation to future ocean acidification.
Fertilization success of an arctic sea urchin species, Strongylocentrotus droebachiensis (O. F. Müller, 1776) under CO2-induced ocean acidificationPublished 15 May 2013 Science Leave a Comment
Tags: biological response, reproduction, echinoderms, laboratory, morphology
Sea urchins as broadcasting spawners, release their gametes into open water for fertilization, thus being particularly vulnerable to ocean acidification. In this study, we assessed the effects of different pH scenarios on fertilization success of Strongylocentrotus droebachiensis, collected at Spitsbergen, Arctic. We achieved acidification by bubbling CO2 into filtered seawater using partial pressures (pCO2) of 180, 380, 980, 1400 and 3000 μatm}. Untreated filtered seawater was used as control. We recorded fertilization rates and diagnosed morphological aberrations after post-fertilization periods of 1 h and 3 h under different exposure conditions in experiments with and without pre-incubation of the eggs prior to fertilization. In parallel, we conducted measurements of intracellular pH changes using BCECF/AM in unfertilized eggs exposed to a range of acidified seawater. We observed increasing rates of polyspermy in relation to higher seawater pCO2, which might be due to failures in the formation of the fertilization envelope. In addition, our experiments showed anomalies in fertilized eggs: incomplete lifting-off of the fertilization envelope and blebs of the hyaline layer. Other drastic malformations consisted of constriction, extrusion, vacuolization or degeneration (observed as a gradient from the cortex to the central region of the cell) of the egg cytoplasm, and irregular cell divisions until 2- to 4-cell stages. The intracellular pH (pHi) decreased significantly from 1400 μatm on. All results indicate a decreasing fertilization success at CO2 concentrations from 1400 μatm upwards. Exposure time to low pH might be a threatening factor for the cellular buffer capacity, viability, and development after fertilization.
Effects of ocean acidification on growth and physiology of Ulva lactuca (Chlorophyta) in a rockpool-scenarioPublished 8 May 2013 Science Leave a Comment
Tags: algae, biological response, growth, laboratory, morphology, North Atlantic, photosynthesis, physiology
Rising atmospheric CO2-concentrations will have severe consequences for a variety of biological processes. We investigated the responses of the green alga Ulva lactuca (Linnaeus) to rising CO2-concentrations in a rockpool scenario. U. lactuca was cultured under aeration with air containing either preindustrial pCO2 (280 μatm) or the pCO2 predicted by the end of the 21st century (700 μatm) for 31 days. We addressed the following question: Will elevated CO2-concentrations affect photosynthesis (net photosynthesis, maximum relative electron transport rate (rETR(max)), maximum quantum yield (Fv/Fm), pigment composition) and growth of U. lactuca in rockpools with limited water exchange? Two phases of the experiment were distinguished: In the initial phase (day 1–4) the Seawater Carbonate System (SWCS) of the culture medium could be adjusted to the selected atmospheric pCO2 condition by continuous aeration with target pCO2 values. In the second phase (day 4–31) the SWCS was largely determined by the metabolism of the growing U. lactuca biomass. In the initial phase, Fv/Fm and rETR(max) were only slightly elevated at high CO2-concentrations, whereas growth was significantly enhanced. After 31 days the Chl a content of the thalli was significantly lower under future conditions and the photosynthesis of thalli grown under preindustrial conditions was not dependent on external carbonic anhydrase. Biomass increased significantly at high CO2-concentrations. At low CO2-concentrations most adult thalli disintegrated between day 14 and 21, whereas at high CO2-concentrations most thalli remained integer until day 31. Thallus disintegration at low CO2-concentrations was mirrored by a drastic decline in seawater dissolved inorganic carbon and HCO3−. Accordingly, the SWCS differed significantly between the treatments. Our results indicated a slight enhancement of photosynthetic performance and significantly elevated growth of U. lactuca at future CO2-concentrations. The accelerated thallus disintegration at high CO2-concentrations under conditions of limited water exchange indicates additional CO2 effects on the life cycle of U. lactuca when living in rockpools.
Tags: abundance, biological response, community composition, fungi, laboratory, molecular biology, North Atlantic, prokaryotes
Anthropogenic CO2 emissions are causing an acidification of the world’s oceans. The consequences for marine organisms and especially heterotrophic bacteria remain under debate, and almost nothing is known concerning marine fungi. Both microbial groups are important players in organic matter decomposition and nutrient cycling, and their pH tolerance is known to be broad in relation to the predicted acidification. So far, ocean acidification effects on marine bacterial communities have mainly been investigated in large-scale mesocosm studies. In these systems, indirect effects mediated through complex food web interactions come into play. Until now, these experiments were not carried out in sufficient replication. In this thesis, we chose an alternative approach and investigated bacterial and fungal communities in highly replicated microcosm experiments (1-1.6 L). The duration of the experiments was four weeks. We incubated the natural microbial community from Helgoland Roads (North Sea) at in situ seawater pH, pH 7.82 and pH 7.67. These pH levels represent the present-day situation and acidification at atmospheric CO2 of 700 or 1000 ppm, projected for the southern North Sea for the year 2100. For the bacterial community, different dilution approaches were used to select for different ecological groups. Seasonality was accounted for by repeating the experiment four times (spring, summer, autumn, winter). In a second experiment repeated in two consecutive years, we investigated direct pH effects on marine fungal communities. We additionally isolated marine yeasts and identified them by Matrix-assisted laser desorption/ionization-time-of-flight mass spectrometry (MALDI-TOF MS) and partial sequencing of the large subunit (LSU) rRNA gene. To reveal changes in community structure, we applied the culture-independent fingerprint method automated ribosomal intergenic spacer analysis (ARISA) for both bacteria and fungi. Bacterial communities were furthermore analyzed by 16S ribosomal amplicon pyrosequencing. Abundances were determined by flow cytometry (bacteria) and colony forming unit counts (fungi). To be able to interpret results comprehensively, we determined the natural variability of the carbonate system at Helgoland Roads over a yearly cycle. We found that from September 2010 to September 2011, pH at Helgoland Roads ranged from 8.06 to 8.43, corresponding to partial pressures of carbon dioxide (pCO2) of 215-526 µatm. The acidification predicted for the year 2100 consequently represents a strong perturbation of the system. Bacterial communities developing in the microcosms were primarily influenced by season and dilution, demonstrating that diverse communities had been generated. We predominantly found pH-dependent shifts in bacterial community structure already at pH 7.82. Groups involved in these shifts were different members of Gammaproteobacteria, Flavobacteriaceae, Rhodobacteraceae, Campylobacteraceae and further less abundant groups. While Rhodobacteraceae were consistently less characteristic for reduced pH, Campylobacteraceae profited from pH reduction. For most other bacterial groups however, pH effects were context-dependent, i.e. dependent on season, dilution or an interaction of effects. Regarding bacterial abundance, no pH effect was found. Fungal community structure was significantly different between both years of the experiment, hinting at inter-annual variability. Shifts in response to pH occurred predominantly only at pH 7.67. In contrast, a strong pH effect was observed on fungal abundance. In comparison to in situ pH, fungal numbers were on average 9 times higher at pH 7.82 and 34 times higher at pH 7.67. Concerning marine yeasts, Leucosporidium scottii, Rhodotorula mucilaginosa and related species, as well as Cryptococcus sp. and Debaromyces hansenii reacted positively to low pH. Our findings demonstrate that already small reductions in pH have direct effects on both bacterial and fungal communities. A tipping point for community shifts appears to be reached earlier for bacteria than for fungi. Regarding bacteria and yeasts, both naturally abundant groups and rare species were affected by pH reductions. The strong increase in fungal numbers at reduced pH suggests that with ocean acidification, marine fungi may reach higher importance in marine biogeochemical cycles and as infectious agents. Using a microcosm approach, a robust analysis of direct ocean acidification effects on marine bacterial and fungal communities was accomplished. Results yield valuable hypotheses to test in future large-scale and long-term studies.