Hot summers raise public awareness of toxic cyanobacterial blooms.

Water quality of eutrophic lakes is threatened by harmful cyanobacterial blooms, which are favored by summer heatwaves and expected to intensify with global warming. Societal demands on surface water for drinking, irrigation and recreation are also highest in summer, especially during dry and warm conditions. Here, we analyzed trends in online searches to investigate how public awareness of cyanobacterial blooms is impacted by temperature in nine different countries over almost twenty years. Our findings reveal large seasonal and interannual variation, with more online searches for harmful cyanobacteria in temperate regions during hot summers. Online searches and media attention increased even more steeply with temperature than the incidence of cyanobacterial blooms, presumably because lakes attract more people during warm weather. Overall, our study indicates that warmer summers not only increase cyanobacterial bloom incidence, but also lead to a pronounced increase of the public awareness of toxic cyanobacterial blooms.

The combined effect of pH and dissolved inorganic carbon concentrations on the physiology of plastidic ciliate Mesodinium rubrum and its cryptophyte prey.

Ocean acidification is caused by rising atmospheric partial pressure of CO2 (pCO2) and involves a lowering of pH combined with increased concentrations of CO2 and dissolved in organic carbon in ocean waters. Many studies investigated the consequences of these combined changes on marine phytoplankton, yet only few attempted to separate the effects of decreased pH and increased pCO2. Moreover, studies typically target photoautotrophic phytoplankton, while little is known of plastidic protists that depend on the ingestion of plastids from their prey. Therefore, we studied the separate and interactive effects of pH and DIC levels on the plastidic ciliate Mesodinium rubrum, which is known to form red tides in coastal waters worldwide. Also, we tested the effects on their prey, which typically are cryptophytes belonging to the Teleaulax/Plagioslemis/Geminigera species complex. These cryptophytes not only serve as food for the ciliate, but also as a supplier of chloroplasts and prey nuclei. We exposed M. rubrum and the two cryptophyte species, T. acuta, T. amphioxeia to different pH (6.8 - 8) and DIC levels (∼ 6.5 - 26 mg C L-1) and assessed their growth and photosynthetic rates, and cellular chlorophyll a and elemental contents. Our findings did not show consistent significant effects across the ranges in pH and/or DIC, except for M. rubrum, for which growth was negatively affected only by the lowest pH of 6.8 combined with lower DIC concentrations. It thus seems that M. rubrum is largely resilient to changes in pH and DIC, and its blooms may not be strongly impacted by the changes in ocean carbonate chemistry projected for the end of the 21st century.

Reconciling contrasting effects of nitrogen on host immunity and pathogen transmission using stoichiometric models.

Hosts rely on the availability of nutrients for growth, and for defense against pathogens. At the same time, changes in host nutrition can alter the dynamics of pathogens that rely on their host for reproduction. For primary producer hosts, enhanced nutrient loads may increase host biomass or pathogen reproduction, promoting faster density-dependent pathogen transmission. However, the effect of elevated nutrients may be reduced if hosts allocate a growth-limiting nutrient to pathogen defense. In canonical disease models, transmission is not a function of nutrient availability. Yet, including nutrient availability is necessary to mechanistically understand the response of infection to changes in the environment. Here, we explore the implications of nutrient-mediated pathogen infectivity and host immunity on infection outcomes. We developed a stoichiometric disease model that explicitly integrates the contrasting dependencies of pathogen infectivity and host immunity on nitrogen (N) and parameterized it for an algal-host system. Our findings reveal dynamic shifts in host biomass build-up, pathogen prevalence, and the force of infection along N supply gradients with N-mediated host infectivity and immunity, compared with a model in which the transmission rate was fixed. We show contrasting responses in pathogen performance with increasing N supply between N-mediated infectivity and N-mediated immunity, revealing an optimum for pathogen transmission at intermediate N supply. This was caused by N limitation of the pathogen at a low N supply and by pathogen suppression via enhanced host immunity at a high N supply. By integrating both nutrient-mediated pathogen infectivity and host immunity into a stoichiometric model, we provide a theoretical framework that is a first step in reconciling the contrasting role nutrients can have on host-pathogen dynamics.

Towards a mechanistic understanding of the impacts of nitrogen deposition on producer-consumer interactions.

Nitrogen (N) deposition has increased substantially since the second half of the 20th century due to human activities. This increase of reactive N into the biosphere has major implications for ecosystem functioning, including primary production, soil and water chemistry and producer community structure and diversity. Increased N deposition is also linked to the decline of insects observed over recent decades. However, we currently lack a mechanistic understanding of the effects of high N deposition on individual fitness, species richness and community structure of both invertebrate and vertebrate consumers. Here, we review the effects of N deposition on producer-consumer interactions, focusing on five existing ecological frameworks: C:N:P ecological stoichiometry, trace element ecological stoichiometry, nutritional geometry, essential micronutrients and allelochemicals. We link reported N deposition-mediated changes in producer quality to life-history strategies and traits of consumers, to gain a mechanistic understanding of the direction of response in consumers. We conclude that high N deposition influences producer quality via eutrophication and acidification pathways. This makes oligotrophic poorly buffered ecosystems most vulnerable to significant changes in producer quality. Changes in producer quality between the reviewed frameworks are often interlinked, complicating predictions of the effects of high N deposition on producer quality. The degree and direction of fitness responses of consumers to changes in producer quality varies among species but can be explained by differences in life-history traits and strategies, particularly those affecting species nutrient intake regulation, mobility, relative growth rate, host-plant specialisation, ontogeny and physiology. To increase our understanding of the effects of N deposition on these complex mechanisms, the inclusion of life-history traits of consumer species in future study designs is pivotal. Based on the reviewed literature, we formulate five hypotheses on the mechanisms underlying the effects of high N deposition on consumers, by linking effects of nutritional ecological frameworks to life-history strategies. Importantly, we expect that N-deposition-mediated changes in producer quality will result in a net decrease in consumer community as well as functional diversity. Moreover, we anticipate an increased risk of outbreak events of a small subset of generalist species, with concomitant declines in a multitude of specialist species. Overall, linking ecological frameworks with consumer life-history strategies provides a mechanistic understanding of the impacts of high N deposition on producer-consumer interactions, which can inform management towards more effective mitigation strategies.

Microfluidic Impedance Cytometry for Single-Cell Particulate Inorganic Carbon:Particulate Organic Carbon Measurements of Calcifying Algae.

Calcifying algae, like coccolithophores, greatly contribute to the oceanic carbon cycle and are therefore of particular interest for ocean carbon models. They play a key role in two processes that are important for the effective CO2 flux: The organic carbon pump (photosynthesis) and the inorganic carbon pump (calcification). The relative contribution of calcification and photosynthesis can be measured in algae by the amount of particulate inorganic carbon (PIC) and particulate organic carbon (POC). A microfluidic impedance cytometer is presented, enabling non-invasive and high-throughput assessment of the calcification state of single coccolithophore cells. Gradual modification of the exoskeleton by acidification results in a strong linear fit (R 2 = 0.98) between the average electrical phase and the PIC:POC ratio of the coccolithophore Emiliania huxleyi 920/9. The effect of different CO2 treatments on the PIC:POC ratio, however, is inconclusive, indicating that there is no strong effect observed for this particular strain. Lower PIC:POC ratios in cultures that grew to higher cell densities are found, which are also recorded with the impedance-based PIC:POC sensor. The development of this new quantification tool for small volumes paves the way for high-throughput analysis while applying multi-variable environmental stressors to support projections of the future marine carbon cycle.

Restoring gradual land-water transitions in a shallow lake improved phytoplankton quantity and quality with cascading effects on zooplankton production.

Land-water transition areas play a significant role in the functioning of aquatic ecosystems. However, anthropogenic pressures are posing severe threats on land-water transition areas, which leads to degradation of the ecological integrity of many lakes worldwide. Enhancing habitat complexity and heterogeneity by restoring land-water transition areas in lake systems is deemed a suitable method to restore lakes bottom-up by stimulating lower trophic levels. Stimulating productivity of lower trophic levels (phytoplankton, zooplankton) generates important food sources for declining higher trophic levels (fish, birds). Here, we study ecosystem restoration project Marker Wadden in Lake Markermeer, The Netherlands. This project involved the construction of a 700-ha archipelago of five islands in a degrading shallow lake, aiming to create additional sheltered land-water transition areas to stimulate food web development from its base by improving phytoplankton quantity and quality. We found that phytoplankton quantity (chlorophyll-a concentration) and quality (inversed carbon:nutrient ratio) in the shallow waters inside the Marker Wadden archipelago were significantly improved, likely due to higher nutrient availabilities, while light availability remained sufficient, compared to the surrounding lake. Higher phytoplankton quantity and quality was positively correlated with zooplankton biomass, which was higher inside the archipelago than in the surrounding lake due to improved trophic transfer efficiency between phytoplankton and zooplankton. We conclude that creating new land-water transition areas can be used to increase light and nutrient availabilities and thereby enhancing primary productivity, which in turn can stimulate higher trophic levels in degrading aquatic ecosystems.

Environmental refuges from disease in host-parasite interactions under global change.

The physiological performance of organisms depends on their environmental context, resulting in performance-response curves along environmental gradients. Parasite performance-response curves are generally expected to be broader than those of their hosts due to shorter generation times and hence faster adaptation. However, certain environmental conditions may limit parasite performance more than that of the host, thereby providing an environmental refuge from disease. Thermal disease refuges have been extensively studied in response to climate warming, but other environmental factors may also provide environmental disease refuges which, in turn, respond to global change. Here, we (1) showcase laboratory and natural examples of refuges from parasites along various environmental gradients, and (2) provide hypotheses on how global environmental change may affect these refuges. We strive to synthesize knowledge on potential environmental disease refuges along different environmental gradients including salinity and nutrients, in both natural and food-production systems. Although scaling up from single host-parasite relationships along one environmental gradient to their interaction outcome in the full complexity of natural environments remains difficult, integrating host and parasite performance-response can serve to formulate testable hypotheses about the variability in parasitism outcomes and the occurrence of environmental disease refuges under current and future environmental conditions.

Comment on "Models predict planned phosphorus load reduction will make Lake Erie more toxic".

Hellweger et al. (Reports, 27 May 2022, pp. 1001) predict that phosphorus limitation will increase concentrations of cyanobacterial toxins in lakes. However, several molecular, physiological, and ecological mechanisms assumed in their models are poorly supported or contradicted by other studies. We conclude that their take-home message that phosphorus load reduction will make Lake Erie more toxic is seriously flawed.

Elemental and biochemical nutrient limitation of zooplankton: A meta-analysis.

Primary consumers in aquatic ecosystems are frequently limited by the quality of their food, often expressed as phytoplankton elemental and biochemical composition. However, the effects of these food quality indicators vary across studies, and we lack an integrated understanding of how elemental (e.g. nitrogen, phosphorus) and biochemical (e.g. fatty acid, sterol) limitations interactively influence aquatic food webs. Here, we present the results of a meta-analysis using >100 experimental studies, confirming that limitation by N, P, fatty acids, and sterols all have significant negative effects on zooplankton performance. However, effects varied by grazer response (growth vs. reproduction), specific manipulation, and across taxa. While P limitation had greater effects on zooplankton growth than fatty acids overall, P and fatty acid limitation had equal effects on reproduction. Furthermore, we show that: nutrient co-limitation in zooplankton is strong; effects of essential fatty acid limitation depend on P availability; indirect effects induced by P limitation exceed direct effects of mineral P limitation; and effects of nutrient amendments using laboratory phytoplankton isolates exceed those using natural field communities. Our meta-analysis reconciles contrasting views about the role of various food quality indicators, and their interactions, for zooplankton performance, and provides a mechanistic understanding of trophic transfer in aquatic environments.

Temperature response of aquatic greenhouse gas emissions differs between dominant plant types.

Greenhouse gas (GHG) emissions from small inland waters are disproportionately large. Climate warming is expected to favor dominance of algae and free-floating plants at the expense of submerged plants. Through different routes these functional plant types may have far-reaching impacts on freshwater GHG emissions in future warmer waters, which are yet unknown. We conducted a 1,000 L mesocosm experiment testing the effects of plant type and warming on GHG emissions from temperate inland waters dominated by either algae, free-floating or submerged plants in controls and warmed (+4 °C) treatments for one year each. Our results show that the effect of experimental warming on GHG fluxes differs between dominance of different functional plant types, mainly by modulating methane ebullition, an often-dominant GHG emission pathway. Specifically, we demonstrate that the response to experimental warming was strongest for free-floating and lowest for submerged plant-dominated systems. Importantly, our results suggest that anticipated shifts in plant type from submerged plants to a dominance of algae or free-floating plants with warming may increase total GHG emissions from shallow waters. This, together with a warming-induced emission response, represents a so far overlooked positive climate feedback. Management strategies aimed at favouring submerged plant dominance may thus substantially mitigate GHG emissions.

Impacts of inorganic nutrients on the physiology of a mixoplanktonic ciliate and its cryptophyte prey.

Many marine planktonic ciliates retain functional chloroplasts from their photosynthetic prey and use them to incorporate inorganic carbon via photosynthesis. While this strategy provides the ciliates with carbon, little is known about their ability to incorporate major dissolved inorganic nutrients, such as nitrogen and phosphorus. Here, we studied how ciliates respond to different concentrations of dissolved inorganic nitrogen and phosphorus. Specifically, we tested the direct and indirect effects of nutrient availability on the ciliate Strombidium cf. basimorphum fed the cryptophyte prey Teleaulax amphioxeia. We assessed responses in the rates of growth, ingestion, photosynthesis, inorganic nutrient uptake, and excretion. Our results show that the prey changed its carbon content depending on the nutrient concentrations. Low inorganic nutrient concentrations increased S. cf. basimorphum growth and prey ingestion. The higher carbon content of the prey under these low nutrient conditions likely supported the growth of the ciliate, while the higher carbon:nutrient stoichiometry of the prey led to the higher ingestion rates. The low carbon content of the prey at high nutrient concentrations resulted in reduced growth of S. cf. basimorphum, which indicates that carbon acquired via photosynthesis in the ciliate cannot compensate for the ingestion of prey with low carbon content. In conclusion, our findings show S. cf. basimorphum is not able to utilize dissolved inorganic nitrogen and phosphorus for growth, and this species seems to be well adapted to exploit its prey when grown at low nutrient conditions.

Impacts of sediment resuspension on phytoplankton biomass production and trophic transfer: Implications for shallow lake restoration.

Wind-induced sediment resuspension in shallow lakes may enhance nutrient availability while reducing light availability for phytoplankton growth, thereby affecting the entire food-web. Lake restoration projects that reduce wind-induced resuspension are expected to enhance trophic transfer efficiencies, thereby improving food-web structure and functioning. Yet, reduced resuspension may also lead to lower nutrient concentrations in the water column, promote benthic algae development, reduce phytoplankton biomass production and thereby reduce secondary production by zooplankton. Lake Markermeer is a shallow delta lake in The Netherlands subject to wind-induced sediment resuspension. Restoration project Marker Wadden consists of newly built islands aiming to reduce sediment resuspension and promote higher trophic levels. Here, we tested the effects of reduced sediment resuspension on phytoplankton biomass build-up, benthic algae development, and zooplankton abundances at different temperatures in a 14-day indoor microcosm experiment. We used Marker Wadden sediment with three resuspension intensities combined with three temperatures, to also test effects of higher temperatures in shallow sheltered waters. Reduced sediment resuspension decreased nutrient concentrations and phytoplankton biomass build-up, while increasing light availability and enhancing benthic algae biomass development. Reduced sediment resuspension furthermore increased zooplankton biomass. Enhanced sediment resuspension and higher temperatures synergistically interacted, maintaining a high level of inorganic suspended solids. Our experimental results are in line with long-term seasonal observations from Lake Markermeer. Our findings demonstrate that for shallow lakes suffering from wind effects, measures such as Marker Wadden aimed at reducing sediment resuspension can be effective in restoring secondary production and supporting higher trophic levels.

Intraspecific variation in multiple trait responses of Alexandrium ostenfeldii towards elevated pCO2.

Dissolved oceanic CO2 concentrations are rising as result of increasing atmospheric partial pressure of CO2 (pCO2), which has large consequences for phytoplankton. To test how higher CO2 availability affects different traits of the toxic dinoflagellate Alexandrium ostenfeldii, we exposed three strains of the same population to 400 and 1,000 µatm CO2, and measured traits including growth rate, cell volume, elemental composition, 13C fractionation, toxin content, and volatile organic compounds (VOCs). Strains largely increased their growth rates and particulate organic carbon and nitrogen production with higher pCO2 and showed significant changes in their VOC profile. One strain showed a significant decrease in both PSP and cyclic imine content and thereby in cellular toxicity. Fractionation against 13C increased in response to elevated pCO2, which may point towards enhanced CO2 acquisition and/or a downscaling of the carbon concentrating mechanisms. Besides consistent responses in some traits, other traits showed large variation in both direction and strength of responses towards elevated pCO2. The observed intraspecific variation in phenotypic plasticity of important functional traits within the same population may help A. ostenfeldii to negate the effects of immediate environmental fluctuations and allow populations to adapt more quickly to changing environments.

Elements of disease in a changing world: modelling feedbacks between infectious disease and ecosystems.

An overlooked effect of ecosystem eutrophication is the potential to alter disease dynamics in primary producers, inducing disease-mediated feedbacks that alter net primary productivity and elemental recycling. Models in disease ecology rarely track organisms past death, yet death from infection can alter important ecosystem processes including elemental recycling rates and nutrient supply to living hosts. In contrast, models in ecosystem ecology rarely track disease dynamics, yet elemental nutrient pools (e.g. nitrogen, phosphorus) can regulate important disease processes including pathogen reproduction and transmission. Thus, both disease and ecosystem ecology stand to grow as fields by exploring questions that arise at their intersection. However, we currently lack a framework explicitly linking these disciplines. We developed a stoichiometric model using elemental currencies to track primary producer biomass (carbon) in vegetation and soil pools, and to track prevalence and the basic reproduction number (R0 ) of a directly transmitted pathogen. This model, parameterised for a deciduous forest, demonstrates that anthropogenic nutrient supply can interact with disease to qualitatively alter both ecosystem and disease dynamics. Using this element-focused approach, we identify knowledge gaps and generate predictions about the impact of anthropogenic nutrient supply rates on infectious disease and feedbacks to ecosystem carbon and nutrient cycling.

Coccolithophore calcification studied by single-cell impedance cytometry: Towards single-cell PIC:POC measurements.

Since the industrial revolution 30% of the anthropogenic CO2 is absorbed by oceans, resulting in ocean acidification, which is a threat to calcifying algae. As a result, there has been profound interest in the study of calcifying algae, because of their important role in the global carbon cycle. The coccolithophore Emiliania huxleyi is considered to be globally the most dominant calcifying algal species, which creates a unique exoskeleton from inorganic calcium carbonate platelets. The PIC (particulate inorganic carbon): POC (particulate organic carbon) ratio describes the relative amount of inorganic carbon in the algae and is a critical parameter in the ocean carbon cycle. In this research we explore the use of microfluidic single-cell impedance spectroscopy in the field of calcifying algae. Microfluidic impedance spectroscopy enables us to characterize single-cell electrical properties in a non-invasive and label-free way. We use the ratio of the impedance at high frequency vs. low frequency, known as opacity, to discriminate between calcified coccolithophores and coccolithophores with a calcite exoskeleton dissolved by acidification (decalcified). We have demonstrated that using opacity we can discriminate between calcified and decalcified coccolithophores with an accuracy of 94.1%. We have observed a correlation between the measured opacity and the cell height in the channel, which is supported by FEM simulations. The difference in cell density between calcified and decalcified cells can explain the difference in cell height and therefore the measured opacity.

Trophic position, elemental ratios and nitrogen transfer in a planktonic host-parasite-consumer food chain including a fungal parasite.

Parasitism is arguably the most commonly occurring consumer strategy. However, only a few food web studies assess how well stable isotopes reflect the trophic position of parasitic consumers and results are variable. Even fewer studies have measured the nutrient transfer by parasitic consumers, hindering an assessment of their role in nutrient transfer through food webs. Here we used a food chain consisting of a diatom as host, a chytrid as its parasitic consumer and a rotifer as the predatory consumer of the chytrid, to assess the trophic position of all three food-chain components using their natural 13C and 15N isotope signatures, and to measure the nitrogen transfer from the host via the chytrid to the rotifer by tracing 15N of a labelled host up the food chain. Additionally, we measured the carbon to nitrogen (C:N) ratios of all food-chain components. Natural isotope abundance results showed no clear 15N enrichment in the chytrid or rotifer relative to the primary producer. However, estimates of nitrogen transfer indicated that about 14% of host nitrogen was transferred per day from host to chytrid during infection epidemics and that some of this nitrogen was also transferred onward to the rotifer. Moreover, C:N ratios decreased with trophic level, suggesting that the chytrid provided a high-quality food source to the rotifer. In conclusion, our results support the "mycoloop". The mycooloop proposes that chytrid infections allow the transfer of nutrients bound in large, inedible phytoplankton to zooplankton through the production of edible transmission spores, thereby rerouting nutrients back into the food web.

Disease-mediated ecosystem services: Pathogens, plants, and people.

Despite the ubiquity of pathogens in ecological systems, their roles in influencing ecosystem services are often overlooked. Pathogens that infect primary producers (i.e., plants, algae, cyanobacteria) can have particularly strong effects because autotrophs are responsible for a wide range of provisioning, regulating, and cultural services. We review the roles of pathogens in mediating ecosystem services provided by autotrophs and outline scenarios in which infection may lead to unexpected outcomes in response to global change. Our synthesis highlights a deficit of information on this topic, and we outline a vision for future research that includes integrative theory and cross-system empirical studies. Ultimately, knowledge about the mediating roles of pathogens on ecosystem services should inform environmental policy and practice.

Effects of Nutrient Limitation on the Synthesis of N-Rich Phytoplankton Toxins: A Meta-Analysis.

Eutrophication has played a major role in the worldwide increase of harmful algal blooms (HABs). Higher input of key nutrients, such as nitrogen (N) and phosphorus (P), can stimulate the growth of harmful algal species in freshwater, estuarine, and coastal marine ecosystems. Some HAB-forming taxa, particularly several cyanobacteria and dinoflagellate species, are harmful through the production of N-rich toxins that have detrimental effects on the environment and human health. Here, we test how changes in nutrient availability affect N-rich toxin synthesis in cyanobacteria and dinoflagellates using a meta-analysis approach. Overall, N-rich toxin content showed an increase with P limitation, while it tended to decrease with N limitation, but we also observed substantial variation in responses both within and across genera and toxin groups. For instance, in response to N limitation, microcystin content varied from a 297% decrease up to a 273% increase, and paralytic shellfish poisoning (PSP) toxin content varied from a 204% decrease to an 82% increase. Cylindrospermopsin, produced by N2-fixing cyanobacteria, showed no clear direction in response to nutrient limitation, and cellular contents of this compound may thus vary independently of nutrient fluctuations. Our results confirm earlier reported stoichiometric regulation of N-rich phytoplankton toxins, showing increased toxin content with an increase in cellular N:P ratios, and vice versa. Thus, changes in N-rich toxin content largely follow the changes in relative cellular N content. Consequently, although nutrient limitation may limit bloom biomass and thereby bloom toxicity, our results warn that P limitation can cause accumulation of cellular toxins and thus lead to unexpected increases in bloom toxicity.

Multiple global change stressor effects on phytoplankton nutrient acquisition in a future ocean.

Predicting the effects of multiple global change stressors on microbial communities remains a challenge because of the complex interactions among those factors. Here, we explore the combined effects of major global change stressors on nutrient acquisition traits in marine phytoplankton. Nutrient limitation constrains phytoplankton production in large parts of the present-day oceans, and is expected to increase owing to climate change, potentially favouring small phytoplankton that are better adapted to oligotrophic conditions. However, other stressors, such as elevated pCO2, rising temperatures and higher light levels, may reduce general metabolic and photosynthetic costs, allowing the reallocation of energy to the acquisition of increasingly limiting nutrients. We propose that this energy reallocation in response to major global change stressors may be more effective in large-celled phytoplankton species and, thus, could indirectly benefit large-more than small-celled phytoplankton, offsetting, at least partially, competitive disadvantages of large cells in a future ocean. Thus, considering the size-dependent responses to multiple stressors may provide a more nuanced understanding of how different microbial groups would fare in the future climate and what effects that would have on ecosystem functioning. This article is part of the theme issue 'Conceptual challenges in microbial community ecology'.

Ocean acidification increases domoic acid contents during a spring to summer succession of coastal phytoplankton.

Enrichment of the oceans with CO2 may be beneficial for some marine phytoplankton, including harmful algae. Numerous laboratory experiments provided valuable insights into the effects of elevated pCO2 on the growth and physiology of harmful algal species, including the production of phycotoxins. Experiments close to natural conditions are the next step to improve predictions, as they consider the complex interplay between biotic and abiotic factors that can confound the direct effects of ocean acidification. We therefore investigated the effect of ocean acidification on the occurrence and abundance of phycotoxins in bulk plankton samples during a long-term mesocosm experiment in the Gullmar Fjord, Sweden, an area frequently experiencing harmful algal blooms. During the experimental period, a total of seven phycotoxin-producing harmful algal genera were identified in the fjord, and in accordance, six toxin classes were detected. However, within the mesocosms, only domoic acid and the corresponding producer Pseudo-nitzschia spp. was observed. Despite high variation within treatments, significantly higher particulate domoic acid contents were measured in the mesocosms with elevated pCO2. Higher particulate domoic acid contents were additionally associated with macronutrient limitation. The risks associated with potentially higher phycotoxin levels in the future ocean warrants attention and should be considered in prospective monitoring strategies for coastal marine waters.