Coastal ecosystems are prone to multiple anthropogenic and natural stressors including eutrophication, acidification, and invasive species. While the growth of some macroalgae can be promoted by excessive nutrient loading and/or elevated pCO2, responses differ among species and ecosystems. Native to the western Pacific Ocean, the filamentous, turf-forming rhodophyte, Dasysiphonia japonica, appeared in estuaries of the northeastern Atlantic Ocean during the 1980s and the northwestern Atlantic Ocean during the late 2000s. Here, we report on the southernmost expansion of the D. japonica in North America and the effects of elevated nutrients and elevated pCO2 on the growth of D. japonica over an annual cycle in Long Island, New York, USA. Growth limitation of the macroalga varied seasonally. During winter and spring, when water temperatures were < 15 °C, growth was significantly enhanced by elevated pCO2 (p < 0.05). During summer and fall, when the water temperature was 15–24 °C, growth was significantly higher under elevated nutrient treatments (p < 0.05). When temperatures reached 28 °C, the macroalga grew poorly and was unaffected by nutrients or pCO2. The δ13C content of regional populations of D. japonica was −30‰, indicating the macroalga is an obligate CO2-user. This result, coupled with significantly increased growth under elevated pCO2 when temperatures were < 15 °C, indicates this macroalga is carbon-limited during colder months, when in situ pCO2 was significantly lower in Long Island estuaries compared to warmer months when estuaries are enriched in metabolically derived CO2. The δ15N content of this macroalga (9‰) indicated it utilized wastewater-derived N and its N limitation during warmer months coincided with lower concentrations of dissolved inorganic N in the water column. Given the stimulatory effect of nutrients on this macroalga and that eutrophication can promote seasonally elevated pCO2, this study suggests that eutrophic estuaries subject to peak annual temperatures < 28 °C may be particularly vulnerable to future invasions of D. japonica as ocean acidification intensifies. Conversely, nutrient reductions would serve as a management approach that would make coastal regions more resilient to invasions by this macroalga.
Predator loss and climate change are hallmarks of the Anthropocene yet their interactive effects are largely unknown. Here, we show that massive calcareous reefs, built slowly by the alga Clathromorphum nereostratum over centuries to millennia, are now declining because of the emerging interplay between these two processes. Such reefs, the structural base of Aleutian kelp forests, are rapidly eroding because of overgrazing by herbivores. Historical reconstructions and experiments reveal that overgrazing was initiated by the loss of sea otters, Enhydra lutris (which gave rise to herbivores capable of causing bioerosion), and then accelerated with ocean warming and acidification (which increased per capita lethal grazing by 34 to 60% compared with preindustrial times). Thus, keystone predators can mediate the ways in which climate effects emerge in nature and the pace with which they alter ecosystems.
Local adaptation can cause predator populations to vary in traits and their effects on prey, but few studies have tested whether divergent predator populations respond differently to acute environmental stressors. We tested how Nucella dogwhelks from 3 populations with natural exposure to distinct environmental regimes in the California Current System altered consumption of mussel prey (Mytilus californianus) in ambient (pH 8.0, 429 µatm partial pressure of CO2 [pCO2]) and acidified (pH 7.6, 1032 µatm pCO2) seawater. Overall, experimental acidification increased the variation in consumption time observed among populations. We found reduced consumption time for the population that experienced more frequent exposure to low pH conditions in nature but not for populations with less prior exposure. Exposure to acidification also altered the individual components of consumption time—search time and handling time—depending on source population. These results indicate that impaired predator performance is not a universal response to acidification, that predation responses to acute acidification can be population specific, and that individual population responses may relate to prior exposure. Our study highlights how population-specific responses to climate change can lead to differences in ecological effects that may restructure prey communities at local scales.
• 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.
Concurrent anthropogenic global climate change and ocean acidification are expected to have a negative impact on calcifying marine organisms. While knowledge of biological responses of organisms to oceanic stress has emerged from single‐species experiments, these do not capture ecologically relevant scenarios where the potential for multi‐organism physiological interactions is assessed. Marine algae provide an interesting case study, as their photosynthetic activity elevates pH in the surrounding microenvironment, potentially buffering more acidic conditions for associated epiphytes. We present findings that indicate increased tolerance of an important epiphytic foraminifera, Marginopora vertebralis , to the effects of increased temperature (±3°C) and p CO2 (~1,000 µatm) when associated with its common algal host, Laurencia intricata . Specimens of M. vertebralis were incubated for 15 days in flow‐through aquaria simulating current and end‐of‐century temperature and pH conditions. Physiological measures of growth (change in wet weight), calcification (measured change in total alkalinity in closed bottles), photochemical efficiency (Fv/Fm ), total chlorophyll, photosynthesis (oxygen flux), and respiration were determined. When incubated in isolation, M. vertebralis exhibited reduced growth in end‐of‐century projections of ocean acidification conditions, while calcification rates were lowest in the high‐temperature, low‐pH treatment. Interestingly, association with L. intricata ameliorated these stress effects with the growth and calcification rates of M. vertebralis being similar to those observed in ambient conditions. Total chlorophyll levels in M. vertebralis decreased when in association with L. intricata , while maximum photochemical efficiency increased in ambient conditions. Net production estimates remained similar between M. vertebralis in isolation and in association with L. intricata , although both production and respiration rates of M. vertebralis were significantly higher when associated with L. intricata . These results indicate that the association with L. intricata increases the resilience of M. vertebralis to climate change stress, providing one of the first examples of physiological buffering by a marine alga that can ameliorate the negative effects of changing ocean conditions.
Larval stages of members of the Abulacraria superphylum including echinoderms and hemichordates have highly alkaline midguts. To date, the reason for the evolution of such extreme pH conditions in the gut of these organisms remains unknown. Here, we test the hypothesis that, analogous to the acidic stomachs of vertebrates, these alkaline conditions may represent a first defensive barrier to protect from environmental pathogens. pH-optimum curves for five different species of marine bacteria demonstrated a rapid decrease in proliferation rates by 50–60% between pH 8.5 and 9.5. Using the marine bacterium Vibrio diazotrophicus, which elicits a coordinated immune response in the larvae of the sea urchin Strongylocentrotus purpuratus, we studied the physiological responses of the midgut pH regulatory machinery to this pathogen. Gastroscopic microelectrode measurements demonstrate a stimulation of midgut alkalization upon infection with V. diazotrophicus accompanied by an upregulation of acid–base transporter transcripts of the midgut. Pharmacological inhibition of midgut alkalization resulted in an increased mortality rate of larvae during Vibrio infection. Reductions in seawater pH resembling ocean acidification conditions lead to moderate reductions in midgut alkalization. However, these reductions in midgut pH do not affect the immune response or resilience of sea urchin larvae to a Vibrio infection under ocean acidification conditions. Our study addressed the evolutionary benefits of the alkaline midgut of Ambulacraria larval stages. The data indicate that alkaline conditions in the gut may serve as a first defensive barrier against environmental pathogens and that this mechanism can compensate for changes in seawater pH.
Bull kelp (Nereocystis leutkeana) forests along the coast for northern California have decreased dramatically as a result of a ‘perfect storm’ of multiple environmental stressors. The disappearance of a predatory sea star and subsequent increase in purple sea urchins (Strongylocentrotus purpuratus) and the recurrence of marine heat waves have caused these once diverse ecosystems to be rapidly converted into relative species-depauperate urchin barrens. By examining the interactive effects of both a rapidly changing abiotic environment and the increase in urchin grazing pressure that is affecting this vital ecosystem, we can better understand its ultimate fate and make better-informed decisions to manage and protect it. As once large and persistent kelp forests are converted into fragmented landscapes of small kelp patches, kelp’s ability to take up dissolved inorganic carbon and reduce nearby acidity and increase both dissolved oxygen and bio-available calcium carbonate may be reduced, preventing it from serving as an environmental stress-free ‘oasis’ of reduced environmental stresses for local marine organisms and affecting ecosystem dynamics. In my first chapter, I examined whether small, fragmented kelp patches are able to retain their ability to alter local seawater chemistry to the same extent a large persistent kelp forests that have been studied previously. I found that in the canopies of small kelp patches, multiple parameters of carbonate chemistry fluctuated more than in the kelp benthos and in adjacent urchin barrens, consistent with metabolic activity by the kelp. Further, kelp fragments increased pH and aragonite saturation and decreased pCO2 during the day to a similar degree as large, intact kelp forests. These results suggest that small kelp patches could mitigate OA stress during the day and serve as spatial and temporal refugia for canopy-dwelling organisms. I also found that the benthic environment in kelp forests and adjacent urchin barrens is subject to unbuffered decreases in temperature, dissolved oxygen and pH. Thus, in chapter two, I assessed how current-day and future-predicted fluctuations in the duration and magnitude of these upwelling-associated stressors would impact the grazing, growth, and survivorship of purple urchins from kelp forest and urchin barren habitats. With upwelling predicted to increase in both intensity and duration with global climate change, understanding whether urchins from different habitats are differentially affected by upwelling-related stressors will give insight into how current and future stressors may be able to help ‘tip the scales’ and convert the increasing number of urchin barrens back into healthy productive kelp forests. I found condition-dependent susceptibility in urchins to increased magnitude and duration upwelling-related stressors. Grazing and gonadal development in kelp forest urchins was most negatively affected by distant future upwelling conditions, whereas in urchin barren urchins, grazing and survival were sensitive to exposure to upwelling in general, and also to increase in magnitudes of acidity, hypoxia, and temperature across both upwelling and non-upwelling events in the future. These results have important implications for population dynamics of urchins and their interactions with bull kelp, which could strongly affect ecosystem dynamics and transitions between kelp forests and urchin barrens. Taken together, the two chapters my thesis provide valuable insight into the potential resilience of bull kelp, a critical foundation species in northeastern Pacific coastal habitats, in the face of a rapidly changing multi-stressor environment.
Brown Ring Disease (BRD) is an infection of the Manila clam Ruditapes philippinarum due to the pathogen Vibrio tapetis . During BRD, clams are facing immunodepression and shell biomineralization alteration. In this paper, we studied the role of pH on the growth of the pathogen and formulated hypothesis on the establishment of BRD by V. tapetis .
Methods and Results
In this study, we monitored the evolution of pH during the growth of V. tapetis in a range of pH and temperatures. We also measured the pH of Manila clam hemolymph and extrapallial fluids during infection by V. tapetis . We highlighted that V. tapetis modulates the external pH during its growth, to a value of 7.70. During the development of BRD, V. tapetis also influences extrapallial fluids and hemolymph pH in vitro in the first hours of exposure and in vivo after 3 days of infection.
Our experiments have shown a close interaction between V. tapetis CECT4600, a pathogen of Manila clam that induces BRD, and the pH of different compartments of the animals during infection. These results indicate that that the bacterium, through a direct mechanism or as a consequence of physiological changes encountered in the animal during infection, is able to interfere with the pH of Manila clam fluids. This pH modification might promote the infection process or at least create an imbalance within the animal, that would favor its persistence. This last hypothesis should be tested in future experiment.
Significance and Impact of Study
This study is the first observation of pH modifications in the context of BRD and might orient future research on the fine mechanisms of pH modulation associated to BRD.
Aquatic ecosystems face a multitude of environmental stressors, including warming and acidification. While warming is expected to have a pronounced effect on plankton communities, many components of the plankton seem fairly robust towards realistic end-of-century acidification conditions. However, interactions of the two stressors and the inclusion of further factors such as nutrient concentration and trophic interactions are expected to change this outcome. We investigated the effects of warming and high CO2 on a nutrient-deplete late summer plankton community from the Kiel Fjord, Baltic Sea, using a mesocosm setup crossing two temperatures with a gradient of CO2. Phytoplankton and microzooplankton (MZP) growth rates as well as biomass, taxonomic composition, and grazing rates of MZP were analysed. We observed effects of high CO2, warming, and their interactions on all measured parameters. The occurrence and direction of the effects were dependent on the phytoplankton or MZP community composition. In addition, the abundance of small-sized phytoplankton was identified as one of the most important factors in shaping the MZP community composition. Overall, our results indicate that an estuarine MZP community used to strong natural fluctuations in CO2 can still be affected by a moderate increase in CO2 if it occurs in combination with warming and during a nutrient-deplete post-bloom situation. This highlights the importance of including trophic interactions and seasonality aspects when assessing climate change effects on marine zooplankton communities.
Numerical models are a suitable tool to quantify impacts of predicted climate change on complex ecosystems but are rarely used to study effects on benthic macroalgal communities. Fucus vesiculosus L. is a habitat‐forming macroalga in the Baltic Sea and alarming shifts from the perennial Fucus community to annual filamentous algae are reported. We developed a box model able to simulate the seasonal growth of the Baltic Fucus–grazer–epiphyte system. This required the implementation of two state variables for Fucus biomass in units of carbon (C) and nitrogen (N). Model equations describe relevant physiological and ecological processes, such as storage of C and N assimilates by Fucus, shading effects of epiphytes or grazing by herbivores on both Fucus and epiphytes, but with species‐specific rates and preferences. Parametrizations of the model equations and required initial conditions were based on measured parameters and process rates in the near‐natural Kiel Outdoor Benthocosm (KOB) experiments during the Biological Impacts of Ocean Acidification project. To validate the model, we compared simulation results with observations in the KOB experiment that lasted from April 2013 until March 2014 under ambient and climate‐change scenarios, that is, increased atmospheric temperature and partial pressure of carbon dioxide. The model reproduced the magnitude and seasonal cycles of Fucus growth and other processes in the KOBs over 1 yr under different scenarios. Now having established the Fucus model, it will be possible to better highlight the actual threat of climate change to the Fucus community in the shallow nearshore waters of the Baltic Sea.
Plankton communities account for at least half of global primary production and play a key role in the global carbon cycle. Warming and acidification may alter the interaction chains in these communities from the bottom and top of the food web. Yet, the relative importance of these potentially complex interactions has not yet been quantified. Here, we examine the isolated and combined effects of warming, acidification, and reductions in phytoplankton and predator abundances in a series of factorial experiments. We find that warming directly impacts the top of the food web, but that the intermediate trophic groups are more strongly influenced by indirect effects mediated by altered top-down interactions. Direct manipulations of predator and phytoplankton abundance reveal similar strong top-down interactions following top predator decline. A meta-analysis of published experiments further supports the conclusion that warming has stronger direct impacts on the top and bottom of the food web rather than the intermediate trophic groups, with important differences between freshwater and marine plankton communities. Our results reveal that the trophic effect of warming cascading down from the top of the plankton food web is a powerful agent of global change.
Coral reefs are declining globally as climate change and local water quality press environmental conditions beyond the physiological tolerances of holobionts—the collective of the host and its microbial symbionts. To assess the relationship between symbiont composition and holobiont stress tolerance, community diversity metrics were quantified for dinoflagellate endosymbionts (Family: Symbiodiniaceae) from eight Acropora millepora genets that thrived under or responded poorly to various stressors. These eight selected genets represent the upper and lower tails of the response distribution of 40 coral genets that were exposed to four stress treatments (and control conditions) in a 10‐day experiment. Specifically, four ‘best performer’ coral genets were analyzed at the end of the experiment because they survived high temperature, high pCO2, bacterial exposure, or combined stressors, whereas four ‘worst performer’ genets were characterized because they experienced substantial mortality under these stressors. At the end of the experiment, seven of eight coral genets mainly hosted Cladocopium symbionts, whereas the eighth genet was dominated by both Cladocopium and Durusdinium symbionts. Symbiodiniaceae alpha and beta diversity were higher in worst performing genets than in best performing genets. Symbiont communities in worst performers also differed more after stress exposure relative to their controls (based on normalized proportional differences in beta diversity), than did best performers. A generalized joint attribute model estimated the influence of host genet and treatment on Symbiodiniaceae community composition and identified strong associations among particular symbionts and host genet performance, as well as weaker associations with treatment. Although dominant symbiont physiology and function contribute to host performance, these findings emphasize the importance of symbiont community diversity and stochasticity as components of host performance. Our findings also suggest that symbiont community diversity metrics may function as indicators of resilience and have potential applications in diverse disciplines from climate change adaptation to agriculture and medicine.
Many microorganisms engaged in host-microbe interactions pendulate between a free-living phase and a host-affiliated stage. How adaptation to stress during the free-living phase affects host-microbe associations is unclear and understudied. To explore this topic, the symbiosis between Hawaiian bobtail squid (Euprymna scolopes) and the luminous bacterium Vibrio fischeri was leveraged for a microbial experimental evolution study. V. fischeri experienced adaptation to extreme pH while apart from the squid host. V. fischeri was serially passaged for 2000 generations to the lower and upper pH growth limits for this microorganism, which were pH 6.0 and 10.0, respectively. V. fischeri was also serially passaged for 2000 generations to vacillating pH 6.0 and 10.0. Evolution to pH stress both facilitated and impaired symbiosis. Microbial evolution to acid stress promoted squid colonization and increased bioluminescence for V. fischeri , while symbiont adaptation to alkaline stress diminished these two traits. Oscillatory selection to acid and alkaline stress also improved symbiosis for V. fischeri , but the facilitating effects were less than that provided by microbial adaptation to acid stress. In summary, microbial adaptation to harsh environments amid the free-living phase may impact the evolution of host-microbe interactions in ways that were not formerly considered.
• Monoculture of oysters produces excess CO2, affecting carbon fluxes.
• Seaweed can eliminate CO2 released by oysters.
• Multi-trophic culture of oysters and seaweed can mitigate oysters monoculture negative impacts.
The physiological responses of aquaculture organisms (e.g., oyster and seaweed) have the potential to affect seawater carbon fluxes and subsequently are affected by these seawater changes. In this study, a laboratory experiment and a field mesocosm experiment were carried out in Daya Bay, southern China. In the laboratory experiment, Portuguese oyster Magallana angulata and the brown seaweed Sargassum hemiphyllum were mono-cultured in 20-L transparent glass bottles for 24 h. Water sample were collected at four incubation time points (i.e. 0 h, 4 h, 12 h and 24 h) to examine their physiological responses across the incubation period. The results showed that the oyster calcification rate was not significantly changed among 4 h, 12 h and 24 h. On the other hand, during the 24 h incubation time, the oyster respiration rate, seawater pH, dissolved oxygen (DO), and CO32– concentration were significantly declined, but the seawater CO2 concentration was increased. For the seaweed, from 0 h to 12 h, seawater CO2 and HCO3– concentrations were significantly declined. However, the seawater pH and DO concentration were increased. In the field experiment, oyster and seaweed were cultured in mesocosm bags. The effects of different culture models of M. angulata and S. hemiphyllum (i.e. oyster monoculture, seaweed monoculture and oyster-seaweed co-culture) on seawater CO2‑carbonate system and air-sea CO2 flux (FCO2) were investigated after 24 h incubation. The results showed that DIC, HCO3– and CO2 concentrations and the partial pressure of CO2 in co-culture bags were significantly lower than the control bags (without any culture organisms) and oyster bags, indicated that S. hemiphyllum can effectively absorb the CO2 released by the oysters. The negative values of air-sea FCO2 in the co-culture bags represent a CO2 sink from the atmosphere to the sea. These results demonstrated that aquaculture organism monoculture could result in a stress for itself, and there could be an interspecies mutual benefit for both M. angulata and S. hemiphyllum in the co-culture system. The negative environmental impacts of mono-trophic oyster aquaculture in this view could be mediated with the multi-trophic inclusion of seaweed.
Since the response to ocean acidification is species specific, differences in responses between predator and prey will alter their interactions, hence affect the population dynamics of both species. Changes in predator prey interactions between a predatory muricid gastropod Reishia clavigera and its prey, the barnacle Amphibalanus amphitrite amphitrite and mussel Brachidontes variabilis under three pCO2 levels (380, 950, and 1250 μatm) were investigated. The searching time for barnacles increased and the ability to locate them decreased at higher pCO2 levels. The movement speed and the prey consumption rate, however, were independent of pCO2. There was no preference towards either B. variabilis or A. amphitrite amphitrite regardless of pCO2. Exposure experiments involving multiple generations are suggested to assess transgenerational effects of ocean acidification and the potential compensation responses before any realistic predictions on the long term changes of population dynamics of the interacting species can be made.
• Increased algal densities reduce photosynthesis and respiration of Ulva conglobata.
• Algal density mediates the interactive effect of increased temperature and lowered pH.
• Altered temperature and pH oppositely affect photosynthetic rate and saturation light.
Growing of macroalgae increases their biomass densities in natural habitats. To explore how the altered algal density impacts their photosynthetic responses to changes of environmental factors, we compared the photosynthesis versus irradiance characteristics of a marine green macroalga Ulva conglobata under low [2.0 g fresh weight (FW) L−1], medium (6.0 g FW L−1) and high biomass densities (12.0 g FW L−1), and under a matrix of temperatures (20, 25, 30 and 35 °C) and pH levels (7.8, 8.2 and 8.6). Increased algal densities decreased the photosynthetic O2 evolution rate among all combined temperature and pH treatments, in parallel with the decrease of light-utilizing efficiency (α, the initial slope) and maximum photosynthetic rate (Pmax) and the increase of light saturation point (EK). Rising temperature interacted with lowered pH to increase the α under low but not under high algal densities. Rising temperature increased the Pmax and decreased the EK under low algal density, but not under high density. Lowered pH promoted the Pmax and EK under all three algal densities. The increased temperature enhanced the dark respiration (Rd) and light compensation point (EC), while the altered pH showed a limited effect. Moreover, the increased algal density reduced the Rd, and had a limited effect on the EC. In addition, our results indicate that changing algal densities caused the complex photophysiological changes in responses to the temperature and pH changes, and these complex responses resolved into a close relation between Rd and Pmax across the matrix of temperatures and pH levels.
• Clams in high pCO2/low pH were more susceptible to infection by pathogenic Vibrios.
• Growth and abundance of Vibrio spp. were greater under high pCO2/low pH.
• Clams reared under high pCO2/low pH seemed to have a broad tolerance range for pH.
• Long-term effect of acidification and susceptibility to vibriosis is understudied.
Ocean acidification alters seawater carbonate chemistry, which can have detrimental impacts for calcifying organisms such as bivalves. This study investigated the physiological cost of resilience to acidification in Mercenaria mercenaria, with a focus on overall immune performance following exposure to Vibrio spp. Larval and juvenile clams reared in seawater with high pCO2 (∼1200 ppm) displayed an enhanced susceptibility to bacterial pathogens. Higher susceptibility to infection in clams grown under acidified conditions was derived from a lower immunity to infection more so than an increase in growth of bacteria under high pCO2. A reciprocal transplant of juvenile clams demonstrated the highest mortality amongst animals transplanted from low pCO2/high pH to high pCO2/low pH conditions and then exposed to bacterial pathogens. Collectively, these results suggest that increased pCO2 will result in immunocompromised larvae and juveniles, which could have complex and pernicious effects on hard clam populations.
• The comprehensive effects of toxic cyanobacteria and high pH on mussels were assessed.
• Interaction between cyanobacteria and high pH on physiological indicator were found.
• Compare to high pH, toxic M. aeruginosa induce more severe oxidative stress response.
• Toxic algae or high pH exposure history showed latent effects on Hyriopsis cumingii.
In lakes and reservoirs, harmful algal blooms and high pH have been deemed to be two important stressors related to eutrophication, especially in the case of CO2 depletion caused by dense blooms. However, the effects of these stressors on the economically important shellfish that inhabit these waters are still not well-understood. This study evaluated the combined effects of the harmful algae Microcystis aeruginosa (0%, 50%, and 100% of total dietary dry weight) and high pH (8.0, 8.5 and 9.0) on the antioxidant responses of the triangle sail mussel H. cumingii. The mussels were exposed to algae and high pH for 14 d, followed by a 7-day depuration period. Reactive oxygen species (ROS) in the mussel hemolymph, antioxidant and detoxifying enzymes, such as glutathione-S-transferase (GST), glutathione (GSH), superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPx), and malondialdehyde (MDA) in the digestive glands were analyzed during the experimental period. GST, SOD and GPx activity levels and the content of GSH increased following exposure to toxic M. aeruginosa, whereas CAT activity was inhibited. pH showed no significant effects on the immune defense mechanisms and detoxification processes. However, a high pH could cause increased ROS and MDA levels, resulting in oxidative injury. After a 7-day depuration period, exposure to toxic M. aeruginosa or high pH resulted in latent effects for most of the examined parameters. The treatment group exposed to the highest pH (9.0) displayed an increased oxidation state compared with the other pH treatments (8.0 and 8.5) for the same concentrations of toxic M. aeruginosa. The trends observed for ROS, MDA, GPx, GST, SOD and GSH levels indicated that a high density of toxic algae could result in severe and continuous effects on mussel health.
• Coculture did not affect growth rate of U. linza but decreased it for S. costatum.
• Elevated CO2 relieved the inhibitory effect of U. linza on growth of S. costatum.
• At elevated CO2, higher temperature increased the growth rate of S. costatum.
• At elevated CO2, higher temperature reduced the growth rate of U. linza.
• Coculture did not affect respiration of U. linza but stimulated it for S. costatum.
Red tide and green tide are two common algal blooms that frequently occur in many areas in the global oceans. The algae causing red tide and green tide often interact with each other in costal ecosystems. However, little is known on how future CO2-induced ocean acidification combined with temperature variation would affect the interaction of red and green tides. In this study, we cultured the red tide alga Skeletonema costatum and the green tide alga Ulva linza under ambient (400 ppm) and future CO2 (1000 ppm) levels and three temperatures (12, 18, 24 °C) in both monoculture and coculture systems. Coculture did not affect the growth rate of U. linza but significantly decreased it for S. costatum. Elevated CO2 relieved the inhibitory effect of U. linza on the growth of S. costatum, particularly for higher temperatures. At elevated CO2, higher temperature increased the growth rate of S. costatum but reduced it for U. linza. Coculture with U. linza reduced the net photosynthetic rate of S. costatum, which was relieved by elevated CO2. This pattern was also found in Chl a content, indicating that U. linza may inhibit growth of S. costatum via harming pigment synthesis and thus photosynthesis. In monoculture, higher temperature did not affect respiration rate of S. costatum but increased it in U. linza. Coculture did not affect respiration of U. linza but stimulated it for S. costatum, which was a signal of responding to biotic and/abiotic stress. The increased growth of S. costatum at higher temperature and decreased inhibition of U. linza on S. costatum at elevated CO2 suggest that red tides may have more advantages over green tides in future warmer and CO2-enriched oceans.
For coral reefs to persist, the rate of CaCO3 production must be greater than the rate of erosion to enable positive growth. Negative impacts of global change (ocean acidiﬁcation and warming) and local stressors (eutrophication, overﬁshing) on accretion co-occur with positive eﬀects of these changes on bioerosion capacity and chemical dissolution by excavating euendolithic organisms. This is especially relevant for reefs characterised with low calcifying rates as they will tip faster into net loss. The Caribbean reefs suﬀered from a decrease by up to 80% in scleractinian coral cover in the past 50 years, their conﬁguration bears very little resemblance with reefs pre1980s, in terms of benthic composition, coral cover and structural complexity. Speciﬁcally, excavating sponges can contribute up to 90% of the total macroborer activity on coral reefs and their rates of bioerosion are positively affected by pCO2. The overarching aim of this thesis was to quantify and understand the accretion and loss terms of coral reef communities with a focus on the interactions of anthropogenic ocean acidiﬁcation and eutrophication with bioerosion by coral-excavating sponges.The use of incubations was central in this piece of work. Changes in the chemical composition of the water overlying sponges and reef communities indicate the relative contribution of metabolic processes such as net calciﬁcation/dissolution and net respiration/production. However, we ﬁrst used ﬂuorescence microscopy to investigate the underlying mechanisms of CaCO3 dissolution by excavating sponges. It revealed that they promote CaCO3 dissolution by decreasing pH at the sponge/coral interface. The high [H+] at this site is achieved through delivery of low-pH vesicles by the etching cells. The enzyme carbonic anhydrase, which is responsible for signiﬁcantly increasing the speed of the reversible reaction H2O+CO2↔H++HCO3−, has been shown to be associated to the sponge’s etching processes and is therefore thought to play a role in the dissolution of CaCO3. By blocking its activity whilst incubating sponges and analysing the rate of dissolution, CA was found to play an important role in speeding up protonation of HCO3− ions at the dissolution site, enabling CO2 to diﬀuse out of the etching area. When exposed to different ranges of ocean acidification and eutrophication, bioerosion rates increased with both variables but no synergistic relation was revealed. Incubations performed at the community level around Saba and Curacao yielded net community calciﬁcation (NCC) rates which were lower than those reported for reef ﬂats worldwide. Still, Saba coral reefs are considered relatively pristine sites compared to the average within the wider Caribbean. Around Curaçao, incubations on reef assemblages dominated by coral yielded even lower NCC rates. Incubations of other benthic assemblages that currently characterized shallow Caribbean reef substrate (such as bioeroding sponges, benthic cyanobacterial mats and sand) all resulted in net dissolution. For both Saba and Curaçao, results suggest that reef calciﬁcation on these sites is barely able to compensate the CaCO3 losses due to dissolution from other opportunistic benthic residents. With the ongoing global and local pressures, the delicate balance between CaCO3 accretion and loss is likely to tip.