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Individual and population level effects of ocean acidification on a predator−prey system with inducible defenses: bryozoan−nudibranch interactions in the Salish Sea

Published 12 December 2018 Science Closed
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Ocean acidification (OA) from in creased oceanic CO2 concentrations imposes significant physiological stresses on many calcifying organisms. OA effects on individual organisms may be synergistically amplified or reduced by inter- and intraspecies interactions as they propagate up to population and community
levels, altering predictions by studies of calcifier responses in isolation. The calcifying colonial bryozoan Membranipora membranacea and the predatory nudibranch Corambe steinbergae comprise a trophic system strongly regulated by predator induced defensive responses and space limitation, presenting a unique system to investigate OA effects on these regulatory mechanisms at individual and population levels. We experimentally quantified OA effects across a range of pH from 7.0 to 7.9 on growth, calcification, senescence and predator-induced spine formation in Membranipora, with or without waterborne predator cue, and on zooid consumption rates in Corambe at Friday Harbor Laboratories, San Juan Island, WA. Membranipora exhibited maximum growth and calcification at moderately low pH (7.6), and continued spine formation in all pH treatments.
Spines reduced Corambe zooid consumption rates, with lower pH weakening this effect. Using a spatially explicit model of colony growth, where colony area
serves as a proxy for colony fitness, we assessed the population-level impacts of these experimentally determined individual-level effects in the context of
space limitation. The area-based fitness costs associated with defense measured at the individual level led to amplified effects predicted for the population level due to competition. Our coupled experimental and modeling results demonstrate the need to consider population-level processes when assessing ecological responses to stresses from changing environments.

Continue reading ‘Individual and population level effects of ocean acidification on a predator−prey system with inducible defenses: bryozoan−nudibranch interactions in the Salish Sea’

The ability of macroalgae to mitigate the negative effects of ocean acidification on four species of North Atlantic bivalve (updated)

Published 25 October 2018 Science Closed
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Coastal ecosystems can experience acidification via upwelling, eutrophication, riverine discharge, and climate change. While the resulting increases in pCO2 can have deleterious effects on calcifying animals, this change in carbonate chemistry may benefit some marine autotrophs. Here, we report on experiments performed with North Atlantic populations of hard clams (Mercenaria mercenaria), eastern oysters (Crassostrea virginica), bay scallops (Argopecten irradians), and blue mussels (Mytilus edulis) grown with and without North Atlantic populations of the green macroalgae, Ulva. In six of seven experiments, exposure to elevated pCO2 levels ( ∼ 1700µatm) resulted in depressed shell- and/or tissue-based growth rates of bivalves compared to control conditions, whereas rates were significantly higher in the presence of Ulva in all experiments. In many cases, the co-exposure to elevated pCO2 levels and Ulva had an antagonistic effect on bivalve growth rates whereby the presence of Ulva under elevated pCO2 levels significantly improved their performance compared to the acidification-only treatment. Saturation states for calcium carbonate (Ω) were significantly higher in the presence of Ulva under both ambient and elevated CO2 delivery rates, and growth rates of bivalves were significantly correlated with Ω in six of seven experiments. Collectively, the results suggest that photosynthesis and/or nitrate assimilation by Ulva increased alkalinity, fostering a carbonate chemistry regime more suitable for optimal growth of calcifying bivalves. This suggests that large natural and/or aquacultured collections of macroalgae in acidified environments could serve as a refuge for calcifying animals that may otherwise be negatively impacted by elevated pCO2 levels and depressed Ω.

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Physiological and biochemical responses of a coralline alga and a sea urchin to climate change: Implications for herbivory

Published 2 October 2018 Science Closed
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Highlights

• Algal metabolism and phenolic content were unaffected by CO2 and temperature treatments.
• CaCO3 content of algae decreased in high CO2 treatments.
• Total sugar content of algae was affected by both CO2 and temperature.
• Sea urchin respiration and feeding increased under high CO2, low temperature.
• Direct effects to sea urchin metabolism drove feeding more than algal palatability.

Abstract

Direct responses to rising temperatures and ocean acidification are increasingly well known for many single species, yet recent reviews have highlighted the need for climate change research to consider a broader range of species, how stressors may interact, and how stressors may affect species interactions. The latter point is important in the context of plant-herbivore interactions, as increasing evidence shows that increasing seawater temperature and/or acidification can alter algal traits that dictate their susceptibility to herbivores, and subsequently, community and ecosystem properties. To better understand how marine rocky shore environments will be affected by a changing ocean, in the present study we investigated the direct effects of short-term, co-occurring increased temperature and ocean acidification on a coralline alga (Jania rubens) and a sea urchin herbivore (Echinometra lucunter) and assessed the indirect effects of these factors on the algal-herbivore interaction. A 21-day mesocosm experiment was conducted with both algae and sea urchins exposed to ambient (24 °C, Low CO2), high-temperature (28 °C, Low CO2), acidified (24 °C, High CO2), or high-temperature plus acidified (28 °C, High CO2) conditions. Algal photosynthesis, respiration, and phenolic content were unaffected by increased temperature and CO2, but calcium carbonate content was reduced under high CO2 treatments in both temperatures, while total sugar content of the algae was reduced under acidified, lower temperature conditions. Metabolic rates of the sea urchin were elevated in the lower temperature, high CO2 treatment, and feeding assays showed that consumption rates also increased in this treatment. Despite some changes to algal chemical composition, it appears that at least under short-term exposure to climate change conditions, direct effects on herbivore metabolism dictated herbivory rates, while indirect effects caused by changes in algal palatability seemed to be of minor importance.

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Comparative effects of seawater acidification on microalgae: Single and multispecies toxicity tests

Published 28 August 2018 Science Closed
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Highlights

• Different responses were observed depending on the species.
• In this work N. gaditana was the most sensitive species to low pH.
• Effects of competence among species were observed in the multispecies control (pH 8.0).
• Effects of competence were eclipsed by the CO2 effects on cultures exposed to pH 6.0.

Abstract

In order to gain knowledge about the potential effects of acidification in aquatic ecosystems, global change research based on microalgae as sentinel species has been often developed. However, these studies are limited to single species tests and there is still a knowledge gap about the behaviour of microalgal communities under this environmental stressor. Thus, the aim of this study was to assess the negative effects of CO2 under an ecologically realistic scenario. To achieve this objective, two types of toxicity tests were developed; i) single toxicity tests and ii) multispecies toxicity tests, in order to evaluate the effects on each species as well as the interspecific competition. For this purpose, three microalgae species (Tetraselmis chuii, Phaeodactylum tricornutum and Nannochloropsis gaditana) were exposed to two selected pH levels (7.4, 6.0) and a control (pH 8.0). The pH values were choosen for testsing different scenarios of CO2 enrichment including the exchange atmosphere-ocean (pH 7.4) and natural or anthropogenic sources of CO2 (pH 6.0). The effects on growth, cell viability, oxidative stress, plus inherent cell properties (size, complexity and autofluorescence) were studied using flow cytometry (FCM). Results showed that T. chuii was the most resistant species to CO2 enrichment with less abrupt changes in terms of cell density, inherent cell properties, oxidative stress and cell viability. Although P. tricornutum was the dominant species in both single and multispecies tests, this species showed a higher decrease in cell density under pH 6.0. Effects of competence were recorded in the multispecies tests (pH 8, control) but this competence was eclipsed by the effects of low pH. The knowledge of biological interactions made by different microalgae species is a useful tool to extrapolate research data from laboratory to the field.

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The bloom-forming macroalgae, Ulva, outcompetes the seagrass, Zostera marina, under high CO2 conditions

Published 23 July 2018 Science Closed
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This study reports on experiments performed with a Northwest Atlantic species of the macroalgae, Ulva, and the seagrass, Zostera marina, grown under ambient and elevated levels of pCO2, and subjected to competition with each other. When grown individually, elevated pCO2 significantly increased growth rates and productivity of Ulva and Zostera, respectively, beyond control treatments (by threefold and 27%, respectively). For both primary producers, significant declines in tissue δ13C signatures suggested that increased growth and productivity were associated with a shift from use of HCO3− toward CO2 use. When grown under higher pCO2, Zostera experienced significant increases in leaf and rhizome carbon content as well as significant increases in leaf carbon-to-nitrogen ratios, while sediments within which high CO2 Zostera were grown had a significantly higher organic carbon content. When grown in the presence of Ulva; however, above- and below-ground productivity and tissue nitrogen content of Zostera were significantly lower, revealing an antagonistic interaction between elevated CO2 and the presence of Ulva. The presence of Zostera had no significant effect on the growth of Ulva. Collectively, this study demonstrates that while Ulva and Zostera can each individually benefit from elevated pCO2 levels, the ability of Ulva to grow more rapidly and inhibit seagrass productivity under elevated pCO2, coupled with accumulation of organic C in sediments, may offset the potential benefits for Zostera within high CO2 environments.

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Interactive effects of ocean acidification with other environmental drivers on marine plankton

Published 16 April 2018 Science Closed
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Planktonic organisms form the base of the marine food web and may be impacted by environmental change in many ways. The interactive effects of multiple, simultaneous climate-driven changes on these organisms are not well understood. This dissertation examined the impacts of ocean acidification in combination with other environmental stressors on marine plankton and determined spatial patterns of one of these potential interactive drivers. Chapter 2 investigated the synergistic effects of ocean acidification and hypoxia on the harmful dinoflagellate Amphidinium carterae. Findings indicated that empirical studies may be crucial to accurately predict organismal responses to multi-stressors. Results also suggested that photorespiration may serve a previously unrecognized role in dinoflagellate metabolism. Chapter 3 examined the combined effects of ocean acidification and lithogenic trace metals on the growth of another harmful dinoflagellate, Cochlodinium polykrikoides. Results indicated that high suspended sediment loads may deliver toxic concentrations of trace elements to marine phytoplankton in acidified coastal ecosystems. Chapter 4 examined the interactive effects of ocean acidification and bacteria on the severity and extent of dissolution in the shells of larval gastropods and the adult pteropod Limacina helicina. Research findings indicated that microbial communities on the shell surfaces of some planktonic molluscs may mediate certain types of shell dissolution in acidified, upwelled waters. Chapter 5 explored the use of thorium isotope fluxes as a proxy for dust and lithogenic iron in the Indian Ocean. Results suggested that the gradient of dust fluxes in the region could impose thresholds for biological productivity. Together, these interdisciplinary studies demonstrate coupled biological and chemical changes in marine ecosystems as a result of increased anthropogenic environmental change.

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The ability of macroalgae to mitigate the negative effects of ocean acidification on four species of North Atlantic bivalve

Published 30 March 2018 Science Closed
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Coastal ecosystems can experience acidification via upwelling, eutrophication, riverine discharge, and climate change. While the resulting increases in pCO2 can have deleterious effects on calcifying animals, this change in carbonate chemistry may benefit some marine autotrophs. Here, we report on experiments performed with North Atlantic populations of hard clams (Mercenaria mercenaria), eastern oysters (Crassostrea virginica), bay scallops (Argopecten irradians), and blue mussels (Mytilus edulis) grown with and without North Atlantic populations of the green macroalgae, Ulva. In 6 of 7 experiments, exposure to elevated pCO2 levels (~ 1,700 µatm) resulted in depressed shell- and/or tissue-based growth rates of bivalves compared to control conditions (p < 0.05) whereas rates were significantly higher in the presence of Ulva in all experiments (p < 0.05). In many cases, the co-exposure elevated pCO2 levels and Ulva had an antagonistic effect on bivalve growth rates whereby the presence of Ulva under elevated pCO2 levels significantly improved their performance compared to the acidification only treatment (p < 0.05). Saturation states for calcium carbonate (Ω) were significantly higher in the presence of Ulva under both ambient and elevated CO2 delivery rates (p < 0.05). Collectively, the results suggest that photosynthesis and/or nitrate assimilation by Ulva increased alkalinity, fostering a carbonate chemistry regime more suitable for optimal growth of calcifying bivalves. This suggests that large natural and/or aquacultured collections of macroalgae in acidified environments could serve as a refuge for calcifying animals that may otherwise be negatively impacted by elevated pCO2 levels and depressed Ω.

Continue reading ‘The ability of macroalgae to mitigate the negative effects of ocean acidification on four species of North Atlantic bivalve’

Abiotic and biotic interactions in the diffusive boundary layer of kelp blades create a potential refuge from ocean acidification

Published 23 March 2018 Science Closed
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  1. Seaweeds are able to modify the chemical environment at their surface, in a micro‐zone called the diffusive boundary layer (DBL), via their metabolic processes controlled by light intensity. Depending on the thickness of the DBL, sessile invertebrates such as calcifying bryozoans or tube‐forming polychaetes living on the surface of the blades can be affected by the chemical variations occurring in this microlayer. Especially in the context of ocean acidification (OA), these microhabitats might be considered as a refuge from lower pH, because during the day photosynthesis temporarily raises the pH to values higher than in the mainstream seawater.
  2. We assessed the thickness and the characteristics of the DBL at two pH levels (today’s average surface ocean pH 8.1 and a reduced pH predicted for the end of the century, pH 7.7) and seawater flows (slow, 0.5 and fast, >8 cm/s) on Ecklonia radiata (kelp) blades. Oxygen and pH profiles from the blade surface to the mainstream seawater were measured with O2 and pH microsensors for both bare blades and blades colonized by the bryozoan Membranipora membranacea.
  3. The DBL was thicker in slow flow compared with fast flow and the presence of bryozoans increased the DBL thickness and shaped the DBL gradient in dark conditions. Net production was increased in the low pH condition, increasing the amount of oxygen in the DBL in both bare and epiphytized blades. This increase drove the daily pH fluctuations at the blade surface, shifting them towards higher values compared with today’s pH. The presence of bryozoans led to lower oxygen concentrations in the DBL and more complex pH fluctuations at the blade surface, particularly at pH 7.7.
  4. Overall, this study, based on microprofiles, shows that, in slow flow, DBL microenvironments at the surface of the kelps may constitute a refuge from OA with pH values higher than those of the mainstream seawater. For calcifying organisms, it could also represent training ground for harsh conditions, with broad daily pH and oxygen fluctuations. These chemical microenvironments, biologically shaped by the macrophytes, are of great interest for the resilience of coastal ecosystems in the context of global change.

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Competitive interactions moderate the effects of elevated temperature and atmospheric CO2 on the health and functioning of oysters

Published 15 November 2017 Science Closed
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Global increases in sea temperatures and atmospheric concentrations of CO2 may affect the health of calcifying shellfish. Little is known, however, about how competitive interactions within and between species may influence how species respond to multiple stressors. We experimentally assessed separate and combined effects of temperature (12 or 16°C) and atmospheric CO2 concentrations (400 and 1000 ppm) on the health and biological functioning of native (Ostrea edulis) and invasive (Crassostrea gigas) oysters held alone and in intraspecific or interspecific mixtures. We found evidence of reduced phagocytosis under elevated CO2 and, when combined with increased temperature, a reduction in the number of circulating haemocytes. Generally, C. gigas showed lower respiration rates relative to O. edulis when the species were in intraspecific or interspecific mixtures. In contrast, O. edulis showed a higher respiration rate relative to C. gigas when held in an interspecific mixture and exhibited lower clearance rates when held in intraspecific or interspecific mixtures. Overall, clearance rates of C. gigas were consistently greater than those of O. edulis. Collectively, our findings indicate that a species’ ability to adapt metabolic processes to environmental conditions can be modified by biotic context and may make some species (here, C. gigas) competitively superior and less vulnerable to future climatic scenarios at local scales. If these conclusions are generic, the relative role of species interactions, and other biotic parameters, in altering the outcomes of climate change will require much greater research emphasis.

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Divergent responses in growth and nutritional quality of coastal macroalgae to the combination of increased pCO2 and nutrients

Published 18 September 2017 Science Closed
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Highlights

  • Growth rates and tissue quality of two common macroalgal species were assessed under conditions of high pCO2 and nutrient loading under monoculture and biculture.
  • Ephemeral macroalgae exhibited significant increases in growth under high pCO2 and high nutrients.
  • Growth rates of perennial macroalgae were unaffected by environmental treatments.
  • Tissue quality of both species increased via decreases in C:N when nutrients were increased.
  • Biculture appears to impact resource acquisition of perennial macroalgae as evidence of higher tissue C:N when compared to monoculture tissue.

Abstract

Coastal ecosystems are subjected to global and local environmental stressors, including increased atmospheric carbon dioxide (CO2) (and subsequent ocean acidification) and nutrient loading. Here, we tested how two common macroalgal species in the Northwest Atlantic (Ulva spp. and Fucus vesiculosus Linneaus) respond to the combination of increased CO2 and nutrient loading. We utilized two levels of pCO2 with two levels of nutrients in a full factorial design, testing the growth rates and tissue quality of Ulva and Fucus grown for 21 days in monoculture and biculture. We found that the opportunistic, fast-growing Ulva exhibited increased growth rates under high pCO2 and high nutrients, with growth rates increasing three-fold above Ulva grown in ambient pCO2 and ambient nutrients. By contrast, Fucus growth rates were not impacted by either environmental factor. Both species exhibited a decline in carbon to nitrogen ratios (C:N) with elevated nutrients, but pCO2 concentration did not alter tissue quality in either species. Species grown in biculture exhibited similar growth rates to those in monoculture conditions, but Fucus C:N increased significantly when grown with Ulva, indicating an effect of the presence of Ulva on Fucus. Our results suggest that the combination of ocean acidification and nutrients will enhance abundance of opportunistic algal species in coastal systems and will likely drive macroalgal community shifts, based on species-specific responses to future conditions.

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Low recruitment due to altered settlement substrata as primary constraint for coral communities under ocean acidification

Published 14 September 2017 Science Closed
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The future of coral reefs under increasing CO2 depends on their capacity to recover from disturbances. To predict the recovery potential of coral communities that are fully acclimatized to elevated CO2, we compared the relative success of coral recruitment and later life stages at two volcanic CO2 seeps and adjacent control sites in Papua New Guinea. Our field experiments showed that the effects of ocean acidification (OA) on coral recruitment rates were up to an order of magnitude greater than the effects on the survival and growth of established corals. Settlement rates, recruit and juvenile densities were best predicted by the presence of crustose coralline algae, as opposed to the direct effects of seawater CO2. Offspring from high CO2 acclimatized parents had similarly impaired settlement rates as offspring from control parents. For most coral taxa, field data showed no evidence of cumulative and compounding detrimental effects of high CO2 on successive life stages, and three taxa showed improved adult performance at high CO2 that compensated for their low recruitment rates. Our data suggest that severely declining capacity for reefs to recover, due to altered settlement substrata and reduced coral recruitment, is likely to become a dominant mechanism of how OA will alter coral reefs.

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Ocean acidification alters zooplankton communities and increases top-down pressure of a cubozoan predator

Published 11 September 2017 Science Closed
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The composition of local ecological communities is determined by the members of the regional community that are able to survive the abiotic and biotic conditions of a local ecosystem. Anthropogenic activities since the industrial revolution have increased atmospheric CO2 concentrations, which have in turn decreased ocean pH and altered carbonate ion concentrations: so called ocean acidification (OA). Single-species experiments have shown how OA can dramatically affect zooplankton development, physiology and skeletal mineralization status, potentially reducing their defensive function and altering their predatory and antipredatory behaviors. This means that increased OA may indirectly alter the biotic conditions by modifying trophic interactions. We investigated how OA affects the impact of a cubozoan predator on their zooplankton prey, predominantly Copepoda, Pleocyemata, Dendrobranchiata, and Amphipoda. Experimental conditions were set at either current (pCO2 370 μatm) or end-of-the-century OA (pCO2 1,100 μatm) scenarios, crossed in an orthogonal experimental design with the presence/absence of the cubozoan predator Carybdea rastoni. The combined effects of exposure to OA and predation by C. rastoni caused greater shifts in community structure, and greater reductions in the abundance of key taxa than would be predicted from combining the effect of each stressor in isolation. Specifically, we show that in the combined presence of OA and a cubozoan predator, populations of the most abundant member of the zooplankton community (calanoid copepods) were reduced 27% more than it would be predicted based on the effects of these stressors in isolation, suggesting that OA increases the susceptibility of plankton to predation. Our results indicate that the ecological consequences of OA may be greater than predicted from single-species experiments, and highlight the need to understand future marine global change from a community perspective.

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The organizing effects of elevated CO2 on competition among estuarine primary producers

Published 25 August 2017 Science Closed
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Fossil fuel combustion, eutrophication, and upwelling introduce excess CO2 into coastal zones. The extent to which marine autotrophs may benefit from elevated CO2 will be a function of their carbon limitation and, among other factors, competition with other primary producers. Here, we report on experiments performed with North Atlantic species of Ulva and Gracilaria grown in situ or exposed to ambient (~400 µatm) and elevated pCO2 (~2500 µatm) and/or subjected to competition with each other and/or with natural plankton assemblages. Elevated pCO2significantly increased the growth rates of Gracilaria and Ulva and yielded significant declines in tissue δ13C, suggesting that increased growth was associated with increased CO2 use relative to HCO3−. Gracilaria growth was unaffected by competition with plankton or Ulva, while Ulva experienced significantly reduced growth when competing with Gracilaria or plankton. Dinoflagellates experienced significantly increased growth when exposed to elevated pCO2 but significantly slower growth when competing with Gracilaria. Elevated carbon-to-nitrogen ratios among macroalgae suggested that competition for nitrogen also shaped interactions among autotrophs, particularly Ulva. While some estuarine autotrophs benefit from elevated pCO2, the benefit can change when direct competition with other primary producers is considered with Gracilaria outcompeting Ulva and dinoflagellates outcompeting diatoms under elevated pCO2.

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Effects of ocean acidification and contact with the brown alga Stypopodium zonale on the settlement and early survival of the coral Porites astreoides

Published 24 August 2017 Science Closed
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To evaluate the effects of ocean acidification (OA) and algal presence on the early life-history stages of corals, we conducted an aquarium study that examined the isolated and combined effects of reduced pH (pH 8.10 vs. 7.85) and contact with the alga Stypopodium zonale on the survival, settlement, and post-settlement growth of larvae from the brooding coral Porites astreoides. Two settlement substrates, biofilmed tiles and the crustose coralline alga (CCA) Hydrolithon boergesenii, were initially incubated for 12 d in separate tanks under a factorial combination of low pH and S. zonale contact, and then subjected to a series of settlement assays. Across both substrate types, S. zonale presence significantly reduced coral settlement. Low pH imposed relatively minor effects; however, there was a significant interaction between pH and S. zonale presence for settlement on the CCA substrate, such that low pH exacerbated the negative effects of S. zonale. Post-settlement growth for 2 wk was unaffected by either S. zonale presence or low pH on either substrate. While our results demonstrate that S. zonale contact likely remains a dominant threat to larval survival and settlement, in certain cases, OA may amplify the negative effects of S. zonale presence, highlighting the need to consider multiple factors in studies aimed at assessing the future health of coral reef ecosystems.

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Macroalgae may mitigate ocean acidification effects on mussel calcification by increasing pH and its fluctuations

Published 15 August 2017 Science Closed
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Ocean acidification (OA) is generally assumed to negatively impact calcification rates of marine organisms. At a local scale however, biological activity of macrophytes may generate pH fluctuations with rates of change that are orders of magnitude larger than the long-term trend predicted for the open ocean. These fluctuations may in turn impact benthic calcifiers in the vicinity. Combining laboratory, mesocosm and field studies, such interactions between OA, the brown alga Fucus vesiculosus, the sea grass Zostera marina and the blue mussel Mytilus edulis were investigated at spatial scales from decimetres to 100s of meters in the western Baltic. Macrophytes increased the overall mean pH of the habitat by up to 0.3 units relative to macrophyte-free, but otherwise similar, habitats and imposed diurnal pH fluctuations with amplitudes ranging from 0.3 to more than 1 pH unit. These amplitudes and their impact on mussel calcification tended to increase with increasing macrophyte biomass to bulk water ratio. At the laboratory and mesocosm scales, biogenic pH fluctuations allowed mussels to maintain calcification even under acidified conditions by shifting most of their calcification activity into the daytime when biogenic fluctuations caused by macrophyte activity offered temporal refuge from OA stress. In natural habitats with a low biomass to water body ratio, the impact of biogenic pH fluctuations on mean calcification rates of M. edulis was less pronounced. Thus, in dense algae or seagrass habitats, macrophytes may mitigate OA impact on mussel calcification by raising mean pH and providing temporal refuge from acidification stress.

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Vulnerability of macronutrients to the concurrent effects of enhanced temperature and atmospheric pCO2 in representative shelf sea sediment habitats

Published 9 August 2017 Science Closed
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Fundamental changes in seawater carbonate chemistry and sea surface temperatures associated with the ocean uptake of anthropogenic CO2 are accelerating, but investigations of the susceptibility of biogeochemical processes to the simultaneous occurrence of multiple components of climate change are uncommon. Here, we quantify how concurrent changes in enhanced temperature and atmospheric pCO2, coupled with an associated shift in macrofaunal community structure and behavior (sediment particle reworking and bioirrigation), modify net carbon and nutrient concentrations (NH4-N, NOx-N, PO4-P) in representative shelf sea sediment habitats (mud, sandy-mud, muddy-sand and sand) of the Celtic Sea. We show that net concentrations of organic carbon, nitrogen and phosphate are, irrespective of sediment type, largely unaffected by a simultaneous increase in temperature and atmospheric pCO2. However, our analyses also reveal that a reduction in macrofaunal species richness and total abundance occurs under future environmental conditions, varies across a gradient of cohesive to non-cohesive sediments, and negatively moderates biogeochemical processes, in particular nitrification. Our findings indicate that future environmental conditions are unlikely to have strong direct effects on biogeochemical processes but, particularly in muddy sands, the abundance, activity, composition and functional role of invertebrate communities are likely to be altered in ways that will be sufficient to regulate the function of the microbial community and the availability of nutrients in shelf sea waters.

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Phytoplankton interactions can alter species response to present and future CO2 concentrations

Published 7 August 2017 Science Closed
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Ocean acidification is a direct consequence of carbon dioxide (CO2) dissolution in seawater and has the potential to impact marine phytoplankton. Although community composition and species interactions may be affected, few studies have taken the latter into account. Here, we assessed how species interactions and competition shape physiological responses by testing monospecific and mixed cultures of (1) the haptophyte Phaeocystis globosa and the chain-forming diatoms Chaetoceros sp. and Asterionellopsis glacialis under present CO2 levels, and (2) Chaetoceros sp. and P. globosa under increasing CO2. The interactions established between the 3 phytoplankton cultures were species- and abundance-dependent. The 2 diatoms did not interact; however, in the presence of P. globosa the growth rates of A. glacialis decreased and those of Chaetoceros sp. increased (depending on a Chaetoceros sp. abundance threshold). Conversely, when Chaetoceros sp. was reasonably abundant, P. globosa was also positively affected (alternating between an abundance/biomass-dependent commensalistic and/or mutualistic interaction). Under enhanced CO2 concentrations, the responses of Chaetoceros sp. and P. globosa mixed cultures were altered, mainly due to Chaetoceros sp. showing a physiological optimum at higher CO2 concentrations than P. globosa. While P. globosa was hindered by increased CO2, Chaetoceros sp. registered augmentation of growth rates, chain length and cellular elemental quotas up to ~750 µatm. Our work emphasizes the role of species interactions when addressing effects of enhanced CO2 on marine phytoplankton. Species-specific response trends to increasing CO2 concentrations revealed significant alterations to species interaction and biomass build-up, which may consequently affect future phytoplankton communities’ composition and dynamics.

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Symbiodinium mitigate the combined effects of hypoxia and acidification on a non-calcifying cnidarian

Published 28 April 2017 Science Closed
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Anthropogenic nutrient inputs enhance microbial respiration within many coastal ecosystems, driving concurrent hypoxia and acidification. During photosynthesis, Symbiodinium spp., the microalgal endosymbionts of cnidarians and other marine phyla, produce O2 and assimilate CO2, and thus potentially mitigate the exposure of the host to these stresses. However, such a role for Symbiodinium remains untested for non-calcifying cnidarians. We therefore contrasted the fitness of symbiotic and aposymbiotic polyps of a model host jellyfish (Cassiopea sp.) under reduced O2 (~2.09mgL−1) and pH (~pH 7.63) scenarios in a full factorial experiment. Host fitness was characterised as asexual reproduction and their ability to regulate internal pH and Symbiodinium performance characterised by maximum photochemical efficiency, chla content, and cell density. Acidification alone resulted in 58% more asexual reproduction of symbiotic polyps than aposymbiotic polyps (and enhanced Symbiodinium cell density) suggesting Cassiopea sp. fitness was enhanced by CO2-stimulated Symbiodinium photosynthetic activity. Indeed, greater CO2 drawdown (elevated pH) was observed within host tissues of symbiotic polyps under acidification regardless of O2 conditions. Hypoxia alone produced 22% fewer polyps than ambient conditions regardless of acidification and symbiont status, suggesting Symbiodinium photosynthetic activity did not mitigate its effects. Combined hypoxia and acidification, however, produced similar numbers of symbiotic polyps compared with aposymbiotic kept under ambient conditions, demonstrating that the presence of Symbiodinium was key for mitigating the combined effects of hypoxia and acidification on asexual reproduction. We hypothesise that this mitigation occurred because of reduced photorespiration under elevated CO2 conditions where increased net O2 production ameliorates oxygen debt. We show that Symbiodinium play an important role in facilitating enhanced fitness of Cassiopea sp. polyps, and perhaps also other non-calcifying cnidarian hosts, to the ubiquitous effects of ocean acidification. Importantly we highlight that symbiotic, non-calcifying cnidarians may be particularly advantaged in productive coastal waters that are subject to simultaneous hypoxia and acidification.

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Cumulative effects of ocean acidification, eutrophication, and competition on the growth of two bloom-forming, estuarine macroalgae

Published 17 March 2017 Science Closed
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While there is a growing interest in understanding how marine life will respond to future ocean acidification, many coastal ecosystems currently experience intense acidification in response to upwelling, riverine discharge, and eutrophication. Such acidification can be inhibitory to calcifying animals, but less is known regarding how non-calcifying macro algae may respond to elevated CO2. Additionally, while the ability of some marine autotrophs to benefit from elevated CO2 over others may result in shifts in community structure, such shifts can also be affected by competition between primary producers. In order to examine what role ocean acidification, eutrophication, and competition plays in the growth of marine macroalgae, a series of experiments were performed during summer through fall 2014 and 2015 with North Atlantic populations of Gracilaria tikvahiae and Ulva rigida that were grown in situ within a mesotrophic estuary (Shinnecock Bay, NY, USA) or exposed to normal and elevated, but environmentally realistic, levels of pCO2 and/or nutrients (nitrogen and phosphorus), as well as being subjected to competition with each other as well as with diatom and dinoflagellate assemblages (2015). Across the 2014 and 2015 experiments, the growth rates of Gracilaria were significantly increased by 70% (2014) and 34% (2015) when exposed to elevated levels of pCO2 (p<0.05). Under the same conditions, the growth rates of Ulva were increased by 30% (2014) and 41% (2015). For nearly all 2014 experiments, Gracilaria was unaffected by nutrient enrichment. In contrast, the growth response of Ulva was more complex as this alga experienced significantly (p<0.05) increased growth rates in response to both elevated pCO2 and nutrients and, in two cases, pCO2 and nutrients interacted to provide synergistically enhanced growth. For the 2015 experiments, growth rates of Gracilaria with or without elevated pCO2 were unaffected by the presence of competing plankton or Ulva. In contrast, growth of Ulva was significantly reduced when grown with Gracilaria (p<0.05) and in several experiments, growth rates of Ulva were found to be significantly reduced when competing with plankton (p<0.05). Dinoflagellates grew significantly faster when exposed to elevated pCO2 (p<0.05) but experienced significantly reduced growth rates grown with Gracilaria (p<0.05). Across all experiments, Gracilaria and Ulva experienced significant declines in tissue δ13C signatures, suggesting that increased growth rates were associated with a shift from use of HCO3- to CO2 use. This shift in carbon use coupled with significantly increased growth in response to elevated pCO2 suggests that photosynthesis of these algae was limited by their inorganic carbon supply. For the 2015 experiments, elevated C:N ratios among macroalgae suggested that competition for N also shaped interactions among autotrophs, particularly for Ulva. Collectively, these study demonstrates that while several types of estuarine autotrophs can benefit from elevated pCO2 levels, their relative benefit can change when direct competition with other primary producers is considered with Gracilaria outcompeting Ulva and dinoflagellates outcompeting diatoms under high pCO2.

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Effects of ocean acidification on the potency of macroalgal allelopathy to a common coral

Published 7 February 2017 Science Closed
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Many coral reefs have phase shifted from coral to macroalgal dominance. Ocean acidification (OA) due to elevated CO2 is hypothesised to advantage macroalgae over corals, contributing to these shifts, but the mechanisms affecting coral-macroalgal interactions under OA are unknown. Here, we show that (i) three common macroalgae are more damaging to a common coral when they compete under CO2 concentrations predicted to occur in 2050 and 2100 than under present-day conditions, (ii) that two macroalgae damage corals via allelopathy, and (iii) that one macroalga is allelopathic under conditions of elevated CO2, but not at ambient levels. Lipid-soluble, surface extracts from the macroalga Canistrocarpus (=Dictyota) cervicornis were significantly more damaging to the coral Acropora intermedia growing in the field if these extracts were from thalli grown under elevated vs ambient concentrations of CO2. Extracts from the macroalgae Chlorodesmis fastigiata and Amansia glomerata were not more potent when grown under elevated CO2. Our results demonstrate increasing OA advantages seaweeds over corals, that algal allelopathy can mediate coral-algal interactions, and that OA may enhance the allelopathy of some macroalgae. Other mechanisms also affect coral-macroalgal interactions under OA, and OA further suppresses the resilience of coral reefs suffering blooms of macroalgae.

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