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Harmful algae bloom (HAB) forecasting has developed rapidly over recent decades, but predicting harmful levels of marine biotoxins in shellfish is still a challenge. New discoveries suggest that predator-prey interactions may be an important driver in the formation of HABs. Key species of harmful algae respond to copepod infochemicals with increased toxin production. In addition, copepods feed selectively on less defended prey, which may further promote harmful taxa. Here we explore if eavesdropping on predator-prey dynamics by monitoring zooplankton can improve HAB forecasting. We first examine an 8-yr time series including copepod biomass, harmful algae cells (Dinophysis spp.), and diarrhetic shellfish toxins in blue mussels (Mytilus edulis) using generalized additive models. Models including copepod biomass more accurately predicted okadaic acid in mussels than phytoplankton alone. We then apply this connection more narrowly by analyzing the specific copepod exudates known to induce toxin production, copepodamides, from the mussels sampled in biotoxin monitoring. Adding copepodamide data from shellfish extracts increased model performance compared to copepod biomass. Results suggest that including grazing effects through copepodamide measurements may provide a cost-efficient way to improve accuracy and lead time for predicting the accumulation of microalgal toxins in shellfish.

Compound-specific isotope analyses (CSIA) and multivariate "isotope fingerprinting" track biosynthetic sources and reveal trophic interactions in food webs. However, CSIA have not been widely applied in the study of marine symbioses. Here, we exposed a reef coral (Montipora capitata) in symbiosis with Symbiodiniaceae algae to experimental treatments (autotrophy, mixotrophy, heterotrophy) to test for trophic shifts and amino acid (AA) sources using paired bulk (δ13C, δ15N) and AA-CSIA (δ13CAA, δ15NAA). Treatments did not influence carbon or nitrogen trophic proxies, thereby not supporting nutritional plasticity. Instead, hosts and symbionts consistently overlapped in essential- and nonessential-δ13CAA (11 of 13 amino acids) and trophic- and source-δ15NAA values (9 of 13 amino acids). Host and symbiont trophic-δ15NAA values positively correlated with a plankton end-member, indicative of trophic connections and dietary sources for trophic-AA nitrogen. However, mass balance of AA-trophic positions (TPGlx-Phe) revealed heterotrophic influences to be highly variable (1-41% heterotrophy). Linear discriminant analysis using M. capitata mean-normalized essential-δ13CAA with previously published values (Pocillopora meandrina) showed similar nutrition isotope fingerprints (Symbiodiniaceae vs. plankton) but revealed species-specific trophic strategies. Montipora capitata and Symbiodiniaceae shared identical AA-fingerprints, whereas P. meandrina was assigned to either symbiont or plankton nutrition. Thus, M. capitata was 100% reliant on symbionts for essential-δ13CAA and demonstrated autotrophic fidelity and contrasts with trophic plasticity reported in P. meandrina. While M. capitata AA may originate from host and/or symbiont biosynthesis, AA carbon is Symbiodiniaceae-derived. Together, AA-CSIA/isotope fingerprinting advances the study of coral trophic plasticity and are powerful tools in the study of marine symbioses.

Factors that regulate planktonic communities under lake ice may be vastly different than the open-water season. However, under-ice food webs in temperate lakes are poorly understood, despite expected changes in light availability, ice cover, and snowfall associated with climate change. We hypothesized that light limitation (bottom-up control) outweighs zooplankton grazing (top-down control) on phytoplankton biovolume and community structure under ice in a north temperate lake. Using in situ under-ice mesocosms, we found that light had stronger effects on phytoplankton abundance than zooplankton, as expected. Specifically, low light limited growth of diatoms, cryptophytes, chrysophytes, and chlorophytes. Zooplankton, however, also significantly affected phytoplankton by decreasing diatoms and cryptophytes, in contrast to the common assumption that zooplankton grazing has negligible effects under ice. Ammonia and soluble reactive phosphorus decreased in high light treatments presumably through uptake by phytoplankton, whereas ammonia and soluble reactive phosphorus increased in high zooplankton treatments, likely through excretion. In situ experimental studies are commonly applied to understand food web dynamics in open-water conditions, but are extremely rare under ice. Our results suggest that changes in the light environment under ice have significant, rapid effects on phytoplankton growth and community structure and that zooplankton may play a more active role in winter food webs than previously thought. Changes in snow and ice dynamics associated with climate change may alter the light environment in ice-covered systems and significantly influence community structure.

Size-spectral approaches quantify the efficiency of energy transfer through food webs, but theory and field studies disagree over how changes in temperature, nutrients, and extreme weather impact on this efficiency. We address this at two scales: via 6 years of weekly sampling of the plankton size spectrum at the Plymouth L4 shelf sea site, and via a new, global-scale, meta-analysis of aquatic size spectra. The time series showed that with summertime nutrient starvation, the energy transfer efficiency from picoplankton to macroplankton decreased (i.e., steepening slopes of the size spectra). This reflected increasing dominance by small cells and their microbial consumers. The extreme storms in winter 2013/2014 caused high metazoan mortality, steep size-spectral slopes, and reduced plankton biomass. However, recovery was within months, demonstrating an inbuilt resilience of the system. Both L4 and our meta-analysis showed steep slopes of normalized size spectra (median −1.11). This reflects much lower values, either of trophic transfer efficiency (3.5%) or predator–prey mass ratio (569), compared to commonly quoted values. Results from the meta-analysis further showed that to represent energy transfer faithfully, size spectra are best constructed in units of carbon mass and not biovolume, and span a mass range of > 107. When this range is covered, both the meta-analysis and time series show a dome-shaped relationship between spectral slopes and plankton biomass, with steepening slopes under increasingly oligotrophic and eutrophic conditions. This suggests that ocean warming could decrease the efficiency of energy transfer through pelagic food webs via indirect effects of increasing stratification and nutrient starvation.

The mixing and stratification present under the ice during winter can have a profound influence on the following summertime hypolimnetic oxygen levels. During winter, plankton rely on updrafts caused by convection to remain in the photic zone in ice-covered lakes, thus there is a crucial link in winter between light levels, under-ice circulation and dissolved oxygen (DO) production. Detailed observations of temperature and oxygen over 3 winters suggest that radiatively driven convection is correlated to oxygen increases in the mixing layer. Both plankton abundance and dissolved oxygen were maximum near the end of the winter before the ice melted. Oxygen became supersaturated by the end of the severe winter of 2015 when the ice cover duration was the longest, whereas DO was slightly below saturation in the warmer winters of 2016 and 2017. After ice-off, the combination of high-frequency measurements through winter and bi-weekly sampling in spring through to summer suggests that decreases in DO started when spring overturn ended and the water column became weakly stratified, which was very close to the timing of when the mean water column temperature first exceeded 4oC. The implication of this work is that the winter oxygen dynamics are important for the hypolimnetic oxygen concentrations when the water column becomes stratified, which in turn sets initial conditions for the degree of any late summer hypoxia.

Trait-based approaches have broadened our understanding of how the composition of ecological communities responds to environmental drivers. This research has mainly focussed on abiotic factors and competition determining the community trait distribution, while effects of trophic interactions on trait dynamics, if considered at all, have been studied for two trophic levels at maximum. However, natural food webs are typically at least tritrophic. This enables indirect interactions of traits and biomasses among multiple trophic levels leading to underexplored effects on food web dynamics. Here, we demonstrate the occurrence of mutual trait adjustment among three trophic levels in a natural plankton food web (Lake Constance) and in a corresponding mathematical model. We found highly recurrent seasonal biomass and trait dynamics, where herbivorous zooplankton increased its size, and thus its ability to counter phytoplankton defense, before phytoplankton defense actually increased. This is contrary to predictions from bitrophic systems where counter-defense of the consumer is a reaction to prey defense. In contrast, counter-defense of carnivores by size adjustment followed the defense of herbivores as expected. By combining observations and model simulations, we show how the reversed trait dynamics at the two lower trophic levels result from a ���trophic biomass���trait cascade��� driven by the carnivores. Trait adjustment between two trophic levels can therefore be altered by biomass or trait changes of adjacent trophic levels. Hence, analyses of only pairwise trait adjustment can be misleading in natural food webs, while multitrophic trait-based approaches capture indirect biomass���trait interactions among multiple trophic levels.

The enzyme carbonic anhydrase (CA) is crucial to many physiological processes involving CO₂, from photosynthesis and respiration, to calcification and CaCO₃ dissolution. We present new measurements of CA activity along a North Pacific transect, on samples from in situ pumps, sediment traps, discreet plankton samples from the ship's underway seawater line, plankton tows, and surface sediment samples from multicores. CA activity is highest in the surface ocean and decreases with depth, both in suspended and sinking particles. Subpolar gyre surface particles exhibit 10× higher CA activity per liter of seawater compared to subtropical gyre surface particles. Activity persists to 4700 m in the subpolar gyre, but only to 1000 m in the subtropics. All sinking CA activity normalized to particulate organic carbon (POC) follows a single relationship (CA/POC = 1.9 ± 0.2 × 10⁻⁷ mol mol⁻¹). This relationship is consistent with CA/POC values in subpolar plankton tow material, suspended particles, and core top sediments. We hypothesize that most subpolar CA activity is associated with rapidly sinking diatom blooms, consistent with a large mat of diatomaceous material identified on the seafloor. Compared to the basin‐wide sinking CA/POC relationship, a lower subtropical CA/POC suggests that the inventory of subtropical biomass is different in composition from exported material. Pteropods also demonstrate substantial CA activity. Scaled to the volume within pteropod shells, first‐order CO₂ hydration rate constants are elevated ≥ 1000× above background. This kinetic enhancement is large enough to catalyze carbonate dissolution within microenvironments, providing observational evidence for CA‐catalyzed, respiration‐driven CaCO₃ dissolution in the shallow North Pacific.

Despite the importance of shallow lakes worldwide, knowledge of microbial components, the base of their food webs, remains scarce. To close this gap, we investigated planktonic microbial food webs, in particular protistan bacterivory (for both ciliates and heterotrophic nanoflagellates [HNF]), in 10 shallow hypertrophic fishponds in South Bohemia (Czech Republic). We used fluorescently labeled bacteria as bacterivory tracers to estimate how abundant protistan populations in fishponds (4–25 × 103 HNF mL−1 and 55–770 ciliates mL−1) contribute to total bacterial mortality. Fluorescence microscopy, innovative image processing tools, and quantitative protargol staining were combined to detect major bacterivorous and omnivorous ciliate taxa. We quantified bacterial production, bacterivory by individual ciliate species, total ciliates, and total protistan bacterivory in all fishponds. On average, ciliate bacterivory was comparable to that of HNF, accounting for 56% and 44% of total protistan grazing, respectively. We found that primarily bacterivorous Peritrichia (genera Vorticella, Epistylis) and Scuticociliata (Cyclidium spp.) contributed only moderately (mean 26%) to total ciliate bacterivory. Unexpectedly, but highly abundant omnivorous Halteria/Pelagohalteria (Stichotrichia) and, to a lesser extent, also omnivorous Rimostrombidium spp. (Oligotrichia) contributed significantly more (mean 71%) to total ciliate bacterivory than typical bacterivorous taxa. This suggests that unselective grazers, which feed on a broader size spectrum from bacteria to small algae, may have a considerable competitive advantage in hypertrophic environments rich in small particles. Moreover, a meta‐analysis of available literature data supports our hypothesis that the role of ciliate bacterivory increases significantly, relative to HNF bacterivory, along a trophic gradient toward hypertrophic habitats.

Plankton are an extremely diverse and polyphyletic group, exhibiting a large range in morphological and physiological traits. Here, we apply automated optical techniques, provided by the pulse‐shape recording automated flow cytometer—CytoSense—to investigate trait variability of phytoplankton and plastidic ciliates in Arctic and Atlantic waters of the subpolar North Atlantic. We used the bio‐optical descriptors derived from the CytoSense (light scattering [forward and sideward] and fluorescence [red, yellow/green and orange from chlorophyll a , degraded pigments, and phycobiliproteins, respectively]) and translated them into functional traits to demonstrate ecological trait variability along an environmental gradient. Cell size was the master trait varying in this study, with large photosynthetic microplankton (> 20 μ m in cell diameter), including diatoms as single cells and chains, as well as plastidic ciliates found in Arctic waters, while small‐sized phytoplankton groups, such as the picoeukaryotes (μ m) and the cyanobacteria Synechococcus were dominant in Atlantic waters. Morphological traits, such as chain/colony formation and structural complexity (i.e., cellular processes, setae, and internal vacuoles), appear to favor buoyancy in highly illuminated and stratified Arctic waters. In Atlantic waters, small cell size and spherical cell shape, in addition to photo‐physiological traits, such as high internal pigmentation, offer chromatic adaptation for survival in the low nutrient and dynamic mixing waters of the Atlantic Ocean. The use of automated techniques that quantify ecological traits holds exciting new opportunities to unravel linkages between the structure and function of plankton communities and marine ecosystems.

Understanding how gelatinous zooplankton communities are structured by local hydrography and physical forcing has important implications for fisheries and higher trophic predators. Although a large body of research has described how fronts, hydrographic boundaries, and different water masses (e.g., mixed vs. stratified) influence phytoplankton and zooplankton communities, comparatively few studies have investigated their influence on gelatinous zooplankton communities. In July 2015, 49 plankton samples were collected from 50 m depth to the surface, across five transects in the Celtic Sea, of which, four crossed the Celtic Sea Front. Two distinct gelatinous communities were found in this dynamic shelf sea: a cold water community in the cooler mixed water that mainly contained neritic taxa and a warm water community in the warmer stratified water that contained a mixture of neritic and oceanic taxa. The gelatinous biomass was 40% greater in the warm water community (∼ 2 mg C m−3) compared with the cold water community (∼ 1.3 mg C m−3). The warm water community was dominated by Aglantha digitale, Lizzia blondina, and Nanomia bijuga, whereas the cold water community was dominated by Clytia hemisphaerica and ctenophores. Physonect siphonophores contributed > 36% to the gelatinous biomass in the warm water community, and their widespread distribution suggests they are ecologically more important than previously thought. A distinct oceanic influence was also recorded in the wider warm water zooplankton community, accounting for a ∼ 20 mg C m−3 increase in biomass in that region.

Temperature sensitivity of plankton in terms of activation energy (Ea, eV) in the Arrhenius equation is critical for predicting how marine productivity and carbon export will respond to ocean warming. In this study, we quantified the temperature responses of phytoplankton growth rate and microzooplankton grazing rate by conducting short-term temperature modulation experiments on natural communities at two subtropical sites with contrasting nutrient conditions. Our results showed that the activation energy of phytoplankton growth rate (Ea = 0.36 eV, 95% confidence interval [CI] = 0.28–0.44 eV) at each station was less than that of microzooplankton grazing rate (Ea = 0.53 eV, 95% CI = 0.47–0.59 eV), indicating an increasing grazing pressure on phytoplankton under warming conditions. Although the difference is consistent with that reported in previous studies, it is very likely to arise from another reason, i.e., differential proximities of the optimal temperature (Topt in nonlinear temperature responses of rates) of phytoplankton and microzooplankton to the environmental temperature, as we found that the environmental temperature is closer to the optimal temperature of phytoplankton growth than to that of microzooplankton grazing in this subtropical environment. Our results suggest that nonlinear temperature responses of plankton should be considered when evaluating and predicting the effects of ocean warming on ecosystem productivity and food web dynamics, especially in subtropical and tropical waters.

Analytical models for the statistical distribution of the gut content of fish larvae in a turbulent ocean environment are compared to data obtained in a field experiment. The proposed model allows the nutrition state and thereby the survival probability of plankton populations to be estimated for given conditions and parameters characterizing their environment, i.e., prey concentrations and turbulence levels. These parameters are all available in the field data. Other parameters such as the capture range and fields of view together with a characteristic time for digesting prey are assumed to be known. The analysis allows an estimate for the probability density of the gut content of plankton in terms of the number of nauplii in the gut. In particular, the analytical results give a basis for evaluating the average gut content of a given plankton population on the basis of basic information concerning the prey concentration and the turbulence intensity. Also analytical models for the prey capture rates are compared with results based on the field data. The analysis emphasizes the effects of turbulence.

17 pages, 6 figures, 2 tables, supporting information https://doi.org/10.1002/lno.11601.-- This is the pre-peer reviewed version of the following article: María Teresa Camarena‐Gómez, Clara Ruiz‐González, Jonna Piiparinen, Tobias Lipsewers, Cristina Sobrino, Ramiro Logares, Kristian Spilling, Bacterioplankton dynamics driven by interannual and spatial variation in diatom and dinoflagellate spring bloom communities in the Baltic Sea, Limnology and Oceanography 66(1): 255-271 (2021), which has been published in final form at https://doi.org/10.1002/lno.11601. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions, In parts of the Baltic Sea, the phytoplankton spring bloom communities, commonly dominated by diatoms, are shifting toward the co‐occurrence of diatoms and dinoflagellates. Although phytoplankton are known to shape the composition and function of associated bacterioplankton communities, the potential bacterial responses to such a decrease of diatoms are unknown. Here we explored the changes in bacterial communities and heterotrophic production during the spring bloom in four consecutive spring blooms across several sub‐basins of the Baltic Sea and related them to changes in environmental variables and in phytoplankton community structure. The taxonomic structure of bacterioplankton assemblages was partially explained by salinity and temperature but also linked to the phytoplankton community. Higher carbon biomass of the diatoms Achnanthes taeniata, Skeletonema marinoi, Thalassiosira levanderi, and Chaetoceros spp. was associated with more diverse bacterial communities dominated by copiotrophic bacteria (Flavobacteriia, Gammaproteobacteria, and Betaproteobacteria) and higher bacterial production. During dinoflagellate dominance, bacterial production was low and bacterial communities were dominated by Alphaproteobacteria, mainly SAR11. Our results suggest that increases in dinoflagellate abundance during the spring bloom will largely affect the structuring and functioning of the associated bacterial communities. This could decrease pelagic remineralization of organic matter and possibly affect the bacterial grazers communities, This study was funded by the Walter and Andrée de Nottbeck Foundation and the Academy of Finland (decision numbers 259164 and 292711). [...] Clara Ruiz González was supported by the GRAMMI project (RTI2018‐099740‐J‐I00, MICIU, Spain). With the funding support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S), of the Spanish Research Agency (AEI)

Littoral benthic primary production is considered the most important energy source of consumers in subarctic lakes. We analyzed essential fatty acid (EFA) and amino acid (EAA) content of 23 littoral benthic macroinvertebrate taxa as well as cladocerans and copepods from pelagic and littoral habitats of 8–9 subarctic lakes to compare their nutritional quality. Pelagic crustacean zooplankton had significantly higher EFA and total FA content (on average 2.6‐fold and 1.6‐fold, respectively) than littoral macroinvertebrates in all our study lakes. Specifically, docosahexaenoic acid (DHA), one of the most important EFA for juvenile fish, was almost exclusively found in pelagic copepods. In littoral macroinvertebrates, only Lymnaea (Gastropoda), Eurycercus (Cladocera), and Gammarus (Amphipoda) contained a low amount of DHA, whereas most littoral invertebrate taxa contained moderate amounts of eicosapentaenoic acid (EPA). The difference in DHA content may explain why so many generalist fish shift their diet to pelagic zooplankton at their peak abundance in mid/late‐summer. Meanwhile, the differences in EAA content between pelagic zooplankton and littoral invertebrates were much lower than for EFA suggesting a wider availability of EAA in subarctic lakes, except for methionine. In the studied subarctic lakes, EFA and EAA variation in consumers was more related to taxon‐specific than lake‐specific characteristics. This indicates that climate‐induced changes in the abundance and community structure of zooplankton vs. littoral macroinvertebrates will be important parameters in determining the availability of EFA and EAA to juvenile fish, and potentially fish production. peerReviewed

WOS:000450233300040; The Plymouth L4 time plankton series in the Western English Channel is a textbook example of a shallow, stratifying shelf sea system. Over its 30 yr of weekly sampling, this site has provided a diverse and contrasting suite of numerical and conceptual models of plankton bloom formation, phenology, and seasonal succession. The most recent of these papers, Kenitz et al. (2017) has initiated this comment, partly because we feel that it has presented a slightly misleading picture of the plankton composition at this site, and of a robust, recurring seasonal succession. We address this by illustrating the extent of inter-annual variability in phenology that occurs at the site, and which needs to be captured better within models. However our main aim is to foster a much better integration of the variety of top-down and bottom-up processes that have all been suggested to be key in driving seasonal succession. Some of these, particularly the multiple grazing and growth controls contributing to the so-called "loophole hypothesis" may be complementary, but others, such as the role of copepod feeding traits in driving species succession (Kenitz et al. 2017) offer testable competing hypotheses. The basic assumptions and outputs of all these models need to be validated more critically, both against time series data and process studies that include the finding of unselective feeding. We suggest that the variability in plankton phenology (and not just mean timing and amplitude) could be used to diagnose the performance of alternative models of plankton succession.

The relationship between zooplankton biomass and phytoplankton biomass can provide insight into the structure and function of lake biological communities. We used a Bayesian network model to analyze a continental-scale data dataset to estimate changes in the relationship between zooplankton (Z) and phytoplankton (P) biomasses along a eutrophication gradient. The Bayesian network model allowed us to combine two different measurements of phytoplankton biomass (chlorophyll a concentration and directly observed biovolume) to improve the precision of estimates of true biomass within each sample. The model also allowed us to estimate separate relationships between P and zooplankton abundance and between P and mean individual zooplankton biomass and then to combine these two relationships into an estimate of seasonal mean zooplankton biomass. The resulting analysis indicated that seasonal mean zooplankton biomass increased proportionally with phytoplankton biomass in oligotrophic lakes, yielding a constant ratio between Z and P and suggested that bottom-up forces determined zooplankton biomass in these systems. In eutrophic lakes, seasonal mean zooplankton biomass was nearly constant with increases in phytoplankton biomass, yielding a decrease in the ratio between Z and P with increasing eutrophication. Bottom-up forces, as quantified by an increase in the proportion of cyanobacteria, accounted for approximately one fifth of the residual variance in the model as the relationship between Z and P changed from direct proportionality in oligotrophic lakes to the nearly constant value of Z observed in eutrophic lakes, suggesting that a combination of both top-down and bottom-up forces likely determined zooplankton biomass in eutrophic lakes.

Lithogenic material such as Saharan dust can be incorporated into organic aggregates and act as ballast, potentially enhancing the marine carbon export via increased sinking velocities of aggregates. We studied the ballasting effects of Saharan dust on the aggregate dynamics in the upwelling region off Cape Blanc (Mauritania). Aggregate formation from a natural plankton community exposed to Saharan dust deposition resulted in higher abundance of aggregates with higher sinking velocities compared to aggregate formation with low dust. This higher aggregate abundance and sinking velocities potentially increased the carbon export 10-fold when the aggregates were ballasted by Saharan dust. After aggregate formation in the surface waters, subsequent sinking through suspended Saharan dust minerals had no influence on aggregate sizes, abundance, and sinking velocities. We found that aggregates formed in the surface ocean off Mauritania were already heavily ballasted with lithogenic material and could therefore not scavenge any additional minerals during their descent. This suggests that carbon export to the deep ocean in regions with high dust deposition is strongly controlled by dust input to the surface ocean while suspended dust particles in deeper water layers do not significantly interact with sinking aggregates.

Elevated allochthonous inputs of organic matter are increasingly recognized as a driver of ecosystem change in lakes, particularly when concurrent with eutrophication. Evaluation of lakes in a nutrient-color paradigm (i.e., based on total phosphorus and true color) enables a more robust approach to research and management. To assess temporal and spatial patterns in nutrient-color status for U.S. lakes and associated food web attributes, we analyzed the U.S. Environmental Protection Agency's National Lakes Assessment (NLA) data. With 1000+ lakes sampled in 2007 and 2012 in a stratified random sampling design, the NLA enables rigorous assessment of lake condition across the continental U.S. We demonstrate that many U.S. lakes are simultaneously experiencing eutrophication and brownification to produce an abundance of "murky" lakes. Overall, "blue" lakes decreased by ~ 18% (46% of lakes in 2007 to 28% in 2012) while "murky" lakes increased by almost 12% (24% of lakes in 2007 to 35.4% in 2012). No statistical differences were observed in the proportions of "green" or "brown" lakes. Regionally, murky lakes significantly increased in the Northern Appalachian, Southern Plains, and Xeric ecoregions. Murky lakes exhibited the highest epilimnetic chlorophyll a concentrations, cyanobacterial densities, and microcystin concentrations. Total zooplankton biomass was also highest in murky lakes, primarily due to increased rotifer and copepod biomass. However, zooplankton : phytoplankton biomass ratios were low, suggesting reduced energy transfer to higher trophic levels. These results emphasize that many lakes in the U.S. are simultaneously "greening" and "browning", with potentially negative consequences for water quality and food web structure.

Diatoms produce a series of cytotoxic secondary metabolites such as polyunsaturated aldehydes (PUA) in response to cell injury or stress. However, little information exists on the PUA effects on the pelagic food web. A set of experiments was conducted in the Chesapeake Bay and the coastal Atlantic waters using dissolved PUA (2E,4E-octadienal and 2E,4E-heptadienal), natural assemblages of phytoplankton and microzooplankton, and the copepod Acartia tonsa. The results demonstrate that PUA primarily acts as deterrent for microzooplankton herbivory on diatoms, while enhancing herbivory on picophytoplankton. This switch should favor PUA-producing diatoms by simultaneously reducing direct grazing losses and competition. Additionally, PUA stimulated copepod predation on microzooplankton, particularly on ciliates. Combined, these effects have the potential to disrupt growth-grazing equilibrium in the pelagic food web and create an opportunity for bloom development.

In a physical model of a thin planktonic layer, the estuarine copepod Acartia tonsa strongly avoided weakly stratified layers of dissolved chemical compounds from the harmful dinoflagellate Karenia brevis. Chemical-induced changes in swimming kinematics allowed copepods to effectively avoid the layer and surrounding volume, highlighting the relevance of harmful alga-grazer interactions at a distance that involve dissolved chemical signals. Avoidance increased significantly with increasing chemical concentration representative of a range of ecologically relevant bloom conditions (1–104 cells/mL equivalent). Under mid to maximal bloom conditions, Acartia displayed visually and hydrodynamically conspicuous avoidance jumps featuring large, rapid displacements with swimming speeds near those reported for predatory escape reactions. Previous findings show K. brevis is both toxic and nutritionally inadequate to A. tonsa when ingested and that exposure to dissolved chemical compounds likely rapidly suppresses effective sampling and grazing behaviors. Thus, copepods have strong incentive to sense and avoid nearby, spatially discrete patches of toxic algae in order to improve fitness by avoiding exposure and/or ingestion and associated negative impacts. Our results suggest harmful alga not only produce deleterious physiological effects in copepod grazers, but chemical-induced behavioral responses also likely alter grazer distributions and top-down control via avoidance reactions (reduced harmful alga-grazer encounter rates). Additionally, predator-prey encounter rates at higher trophic levels are likely enhanced via significant changes in copepod swimming kinematics. These combined mechanisms could protect and sustain harmful blooms contained in subsurface thin layers until blooms reach critical mass and produce widespread impacts at the ecosystem level, the “cryptic bloom” effect.

Highly productive kelps release abundant particulate organic matter into the nearshore environment due to their constant fragmentation and erosion by ocean waves. The contribution of kelp detritus to coastal planktonic food webs has not previously been examined. Here, we demonstrate that detritus derived from a dominant kelp in the Northeast Pacific, Nereocystis luetkeana, provides high-quality food for planktonic sea urchin larvae. Our findings challenge the paradigm that phytoplankton are the main diet for zooplankton in nearshore regions, with implications for modeling of ocean production. Furthermore, at the benthic adult stage, sea urchins can destructively graze kelps causing the kelp ecosystem to collapse; hence, our results have implications for understanding feedback mechanisms that may determine the resilience of kelp ecosystems.

Appendicularians are planktonic marine tunicates with elaborate filter-feeding houses that can efficiently trap particles as small as 0.2 μm. While marine viruses are seldom considered outside their role in disease transmission, we conducted a controlled laboratory experiment to determine if the appendicularian Oikopleura dioica can trap and ingest the Emiliania huxleyi virus (EhV; 160–180 nm diameter). Removal and retention of EhV during 2.5 h and overnight incubations at 15°C were measured using flow cytometry and quantitative polymerase chain reaction specific for the mcp gene of EhV. The fate of retained EhV was tested by quantifying EhV DNA in three biological compartments: house-trapping, ingestion/digestion, and defecation. Clearance rates for EhV varied from approximately 2 mL ind−1 d−1 to 50 mL ind−1 d−1, with highest rates for 4–5 d-old animals. EhV particles were cleared by O. dioica at rates similar to those reported for larger food particles, with mean clearance rates in the 2.5 h incubations ranging from approximately 2 mL ind−1 d−1 to 50 mL ind−1 d−1. This demonstrates efficient virus removal by O. dioica and a previously overlooked link between the microbial loop and the classical marine food web. EhV DNA was readily detectable above background levels in O. dioica houses, gut contents, and faecal pellets, suggesting that appendicularian houses and faecal pellets may contribute to the dispersal of viruses. Furthermore, clearance of EhV and presumably other viruses by O. dioica may be a significant sink for viruses and thus an important factor in regulating the population dynamics of viruses and their hosts.

New resting cyst production is crucial for the survival of many microbial eukaryotes including phytoplankton that cause harmful algal blooms. Production in situ has previously been estimated through sediment trap deployments, but here was instead assessed through estimation of the total number of planktonic cells and new resting cysts produced by a localized, inshore bloom of Alexandrium catenella, a dinoflagellate that is a globally important cause of paralytic shellfish poisoning. Our approach utilizes high frequency, automated water monitoring, weekly observation of new cyst production, and pre- and post-bloom spatial surveys of total resting cyst abundance. Through this approach, new cyst recruitment within the study area was shown to account for at least 10.9% ± 2.6% (SE) of the bloom's decline, ∼ 5× greater than reported from comparable, sediment trap based studies. The observed distribution and timing of new cyst recruitment indicate that: (1) planozygotes, the immediate precursor to cysts in the life cycle, migrate nearer to the water surface than other planktonic stages and (2) encystment occurs after planozygote settlement on bottom sediments. Near surface localization by planozygotes explains the ephemerality of red surface water discoloration by A. catenella blooms, and also enhances the dispersal of new cysts. Following settlement, bioturbation and perhaps active swimming promote sediment infiltration by planozygotes, reducing the extent of cyst redistribution between blooms. The concerted nature of bloom sexual induction, especially in the context of an observed upper limit to A. catenella bloom intensities and heightened susceptibility of planozygotes to the parasite Amoebophrya, is also discussed.

Can we predict long-term trends of lake surface temperature based on air temperature alone? We explore this question by analyzing the performance of a hybrid model (air2water) as a predictive tool for defining scenarios of lake surface temperature in the framework of climate change studies. Employing Lake Tahoe (U.S.A.) as a case study, we apply the model using different air temperature datasets (in situ measurements, gridded observations, and downscaled General Circulation Models). Through a data-driven calibration of the model parameters based on surface water temperature records, we show that air2water provides good performance (root mean square error ∼ 0.5°C, on a monthly scale) regardless of the input dataset. The model is able to accurately capture the historical long-term trend and interannual fluctuations over decades (from 1969 to present), using only 7 yr of monthly measurements of surface water temperature for calibration. Additionally, when used to predict future surface water temperature of the lake, air2water produces the same projections irrespective of the air temperature dataset used to drive the model. This is certainly desirable, but not immediately expected when using a relatively simple model. Overall, the results suggest the high potential and robustness of air2water as a predictive tool for climate change assessment. Lake surface temperature warming of up to 1.1°C (RCP 4.5) and 2.9°C (RCP 8.5) was simulated at the end of the 21st century during summer months in Lake Tahoe. Such a scenario, if realized, would lead to serious consequences on lake water chemistry, primary productivity, plankton community structure, and nutrient cycling.

Acetaldehyde, methanol, acetone, and isoprene are important reactive volatile organic compounds (VOCs) in the oceans that partition to the atmosphere in significant amounts. Reports of potentially high rates of VOC turnover in the North Atlantic suggested that both biotic and abiotic processes are involved. The biological basis for VOC cycling by ocean plankton is unknown, but is potentially important because of VOC contributions to carbon cycle budgets and atmospheric chemistry. We designed dynamic stripping chambers that coupled to a proton transfer reaction mass spectrometer to measure VOC production and consumption by cultured phytoplankton and bacterioplankton. The diatom, Thalassiosira pseudonana, produced acetaldehyde in a light-dependent manner. Acetaldehyde was oxidized by the chemoheterotrophic bacterium, Pelagibacter, at rates that suggest that most acetaldehyde is recycled in the ocean before escaping to the atmosphere. These results show that field observations of acetaldehyde turnover reported previously could be explained by biological activity. Rates of production by phytoplankton cultures of methanol, acetone, and isoprene were also measured. These findings support the conclusion that VOCs are a conduit for carbon transfer directly from phytoplankton to bacterioplankton, with the remainder available for escape to the atmosphere.

Sponges are commonly divided into high (HMA) and low (LMA) microbial-abundance species according to the bacterial biomass in their tissue. These two groups reflect distinct aquiferous structures and feeding strategies. In the NW Mediterranean coralligenous community, HMA and LMA sponges are often packed in dense, multispecies assemblages that cover many pinnacles and overhangs. We investigated the metabolism of HMA and LMA species that cohabitate the coralligenous community by sampling in situ the inhaled and exhaled water. Sponges consumed plankton, dissolved organic carbon (DOC), and ammonium in relation to their abundance in ambient water. The plankton retention efficiency was high for all species. DOC was the main source of C for the sponge species, accounting for ∼ 90% of the examined sources. Nitrogen fluxes markedly differed between the two groups: plankton was the main source of nitrogen for LMAs that excreted dissolved organic nitrogen (DON) and ammonium. The nitrogenous waste products of LMAs were found to be the major source of nitrogen (up to 97%) for HMAs that efficiently removed DON and ammonium and excreted nitrate. The different capacity of both sponge strategies to use dissolved resources suggests a partial trophic niche separation related to HMA-LMA dichotomy as a mechanism facilitating their dense co-existence in the community. Our findings suggest that a mixed assemblage of sponges (and their associated microbes) is able to utilize the suspended particulate and dissolved material more efficiently than a single species population and may contribute to the understanding of the phenomena of the stability and diversity of dense sponge assemblages in oligotrophic habitats.

Size structure of phytoplankton determines to a large degree the trophic interactions in oceanic and coastal waters and eventually the destiny of its biomass. Although, tropical estuarine systems are some of the most productive systems worldwide compared to temperate systems, little is known about phytoplankton biomass size fractions, their contribution to net metabolism, or the ecological factors driving phytoplankton size distribution in tropical estuaries. Hence, we measured the size-fractionated biomass and net metabolism of the plankton community along a salinity and nutrient gradient in the Gulf of Nicoya estuary (Costa Rica), during the dry season. Respiration (23.6 mmol O2 m−3 h−1) was highest at the estuary head, whereas maximum net primary production (23.1 mmol O2 m−3 h−1) was observed in the middle of the estuary, coinciding with the chlorophyll a maximum (15.9 mg m−3). Thus, only the middle section of the estuary was net autotrophic (2.9 g C m−2 d−1), with the rest of the estuary being net heterotrophic. Regression analysis identified light availability, and not nutrients, as the principal factor limiting primary production in the estuary due to increased turbidity. The changes in net metabolism along the estuary were also reflected in the phytoplankton's size structure. Although micro- and picophytoplankton were the most productive fractions overall, in the middle section of the estuary nanophytoplankton dominated primary production, chlorophyll, and autotrophic biomass.

The surface water of the Western Philippine Sea (WPS) receives a significant amount of anthropogenic aerosols from East Asia, serving as an ideal location to investigate the impact of anthropogenic aerosol deposition on trace metal composition and cycling in the Northwestern Pacific Ocean (NWPO). As part of the Taiwan GEOTRACES process study, we have collected size-fractionated plankton from surface water to investigate their metal composition in the open ocean. Elemental ratios in plankton, P- and Al-normalized, are used to evaluate the sources of trace metals and the relative contribution of different metal components in plankton assemblages. Most of the trace metal quotas in plankton are one to two orders of magnitude higher than their intracellular plankton quota, indicating that the majority of the metals are most likely to be extracellularly adsorbed or aggregated on plankton. The metal to Al ratios for most of the trace metals in plankton are also one to two orders of magnitude higher than their lithogenic composition, but are relatively close to metal composition in aerosols collected in situ. This supports the notion that particulate trace metals associated with plankton mainly originate from anthropogenic aerosols, not lithogenic particles. Compared to plankton metal quotas obtained in other oceanic regions, trace metal quotas observed in the WPS rank among the highest for Fe, Mn, Zn, and Cu globally. Our study demonstrates that anthropogenic aerosol deposition has significantly elevated trace metal concentrations in the size-fractionated plankton in the surface water of the NWPO relative to the biological requirements.

Predation is an important source of mortality in zooplankton but factors governing predation risk in marine food webs are still not well understood. Here, we examine the role of zooplankton behavior in determining predation risk. We first quantified motility of copepods with different feeding behaviors (ambush feeding, cruising, and feeding-current feeding). Second, we estimated remote predator detection and escape characteristics of the studied copepods. Third, we proposed a simple behavior-dependent encounter model to predict copepod predation risk from rheotactic predators. Finally, we compared our predictions with predation risk previously determined experimentally. For similar sized copepods, predicted predation risks were similar between feeding-current feeders and cruising feeders, whereas predation was up to 8.5 times lower (range: 1.5–8.5) for ambush feeders. Predicted predation risks further differed between males and females depending on feeding behavior: in ambush feeders males actively search for non-motile females and their predation risk was up to 6 times higher (range 1.2–6) than for females. In contrast, feeding current- and cruising feeders showed small differences in predation risk between genders. In all cases, predicted relative predation risks between particular behaviors were confirmed by empirical data from previous predation experiments. Our results demonstrate that prey behavior of zooplankton may lead to a predictable variation in predation risk from rheotactic predators of up to an order of magnitude, and therefore that individual behavior is an important factor in structuring zooplankton communities.

Rising temperatures in the Arctic Ocean are causing sea ice and glaciers to melt at record breaking rates, which has consequences for carbon cycling in the Arctic Ocean that are yet to be fully understood. Microbial carbon cycling is driven by internal processing of in situ produced organic carbon (OC), however recent research suggests that melt water from sea ice and glaciers could introduce an allochthonous source of OC to the microbial food web with ramifications for the metabolic balance of plankton communities. In this study, we characterized autochthonous and allochthonous sources of OC to the Western Svalbard fjord system using stable isotopes of carbon. We quantified δ13C of eukaryotic and prokaryotic planktonic groups using polar lipid-derived fatty acids as biomarkers in addition to measuring δ13C of marine particulate OC and dissolved OC from glacial runoff. δ13C of bacteria (−22.5‰) was higher than that of glacial runoff OC (−28.5‰) and other phytoplankton groups (−24.7 to −29.1‰), which suggests that marine bacteria preferentially use a third source of OC. We present a Bayesian three-source δ13C mixing model whereby ∼ 60% of bacteria carbon is derived from OC in sea ice, and the remaining carbon is derived from autochthonous production and glacial-derived OC. These results suggest that subsidies of OC from melting glaciers will not likely influence microbial carbon cycling in Svalbard fjords in the future and that further research is needed to determine the effects of melting sea ice on microbial carbon cycling in fjord systems and elsewhere in the Arctic Ocean.

Source at http://dx.doi.org/10.1002/lno.10519 Zooplankton vertical migration enhances the efficiency of the ocean biological pump by translocating carbon (C) and nitrogen (N) below the mixed layer through respiration and excretion at depth. We measured C and N active transport due to diel vertical migration (DVM) in a Svalbard fjord at 79°N. Multifrequency analysis of backscatter data from an Acoustic Zooplankton Fish Profiler moored from January to September 2014, combined with plankton net data, showed that Thysanoessa spp. euphausiids made up > 90% of the diel migrant biomass. Classical synchronous DVM occurred before and after the phytoplankton bloom, leading to a mismatch with intensive primary production during the midnight sun. Zooplankton DVM resulted in C respiration of 0.9 g m−2 and ammonium excretion of 0.18 g N m−2 below 82 m depth between February and April, and 0.2 g C m−2 and 0.04 g N m−2 from 11 August to 9 September, representing > 25% and > 33% of sinking flux of particulate organic carbon and nitrogen, respectively. Such contribution of DVM active transport to the biological pump in this high-Arctic location is consistent with previous measurements in several equatorial to subarctic oceanic systems of the World Ocean. Climate warming is expected to result in tighter coupling between DVM and bloom periods, stronger stratification of the Barents Sea, and northward advection of boreal euphausiids. This may increase the role of DVM in the functioning of the biological pump on the Atlantic side of the Arctic Ocean, particularly where euphausiids are or will be prevalent in the zooplankton community.

While considerable effort has been devoted to understanding the factors regulating the development of phytoplankton blooms, the mechanisms leading to bloom decline and termination have received less attention. Grazing and sedimentation have been invoked as the main routes for the loss of phytoplankton biomass, and more recently, viral lysis, parasitism, and programmed cell death (PCD) have been recognized as additional removal factors. Despite the importance of bloom declines to phytoplankton dynamics, the incidence and significance of various loss factors in regulating phytoplankton populations have not been widely characterized in natural blooms. To understand mechanisms controlling bloom decline, we studied two independent, inshore blooms of Alexandrium fundyense, paying special attention to cell mortality as a loss pathway. We observed increases in the number of dead cells with PCD features after the peak of both blooms, demonstrating a role for cell mortality in their terminations. In both blooms, sexual cyst formation appears to have been the dominant process leading to bloom termination, as both blooms were dominated by small-sized gamete cells near their peaks. Cell death and parasitism became more significant as sources of cell loss several days after the onset of bloom decline. Our findings show two distinct phases of bloom decline, characterized by sexual fusion as the initial dominant cell removal processes followed by elimination of remaining cells by cell death and parasitism.

While decreasing occurrence and duration of lake ice cover is well-documented, biogeochemical dynamics in frozen lakes remain poorly understood. Here, we interpret winter physical and biogeochemical time series from eutrophic Missisquoi Bay (MB) and hyper-eutrophic Shelburne Pond (SP) to describe variable drivers of under ice biogeochemistry in systems of fundamentally different lake-watershed physical configurations (lake area, lake : watershed area). The continuous cold of the 2015 winter drove the MB sediment-water interface to the most severe and persistent suboxic state ever documented at this site, promoting the depletion of redox-sensitive phases in sediments, and an expanding zone of bottom water enriched in reactive species of Mn, Fe, and P. In this context, lake sediment and water column inventories of reactive chemical species were sensitive to the severity and persistence of subfreezing temperatures. During thaws, event provenance and severity impact lake thermal structure and mixing, water column enrichment in P and Fe, and thaw capability to suppress redox front position and internal chemical loading. Nearly identical winter weather manifest differently in nearby SP, where the small surface and watershed areas promoted a warmer, less stratified water column and active phytoplankton populations, impacting biogeochemical dynamics. In SP, Fe and P behavior under ice were decoupled due to active biological cycling, and thaw impacts were different in distribution and composition due to SP's physical structure and related antecedent conditions. We find that under ice biogeochemistry is highly dynamic in both time and space and sensitive to a variety of drivers impacted by climate change.

The seasonal forcing of pelagic communities invokes a succession of the dominant phytoplankton and zooplankton species. Here, we characterize the seasonal succession of the plankton traits and their interactions using observations and model simulations of the plankton community in the western English Channel. We focus on activity traits that characterize the defensive and feeding abilities of zooplankton and distinguish between low risk, low return ambush feeders and high risk, high return feeding-current feeders. While the phytoplankton succession depends on traits related to nutrient acquisition and photosynthesis, it also depends on grazing which couples feeding and motility traits across trophic guilds. Despite interannual variations in the species dominating the protist plankton community, the seasonal trait distribution reveals robust and repeatable seasonal patterns, changing between non-motile cells flourishing in spring and motile community dominating during summer. The zooplankton community is dominated by active feeding-current feeders with peak biomass in the late spring declining during summer. The model reveals how zooplankton grazing reinforces protist plankton seasonal succession and shows how the physical environment controls the vertical structure of plankton communities, where ambush feeders exhibit a preference for greater depths during summer. We characterize the seasonal succession as trophic trait coupling and conjecture that this coupling leads to a trophic trait cascade where successive trophic levels alternate in their expression of activity traits further up in the food chain.

Symbiosis with micro-algae (photosymbiosis) is a common feature among marine planktonic protists, but very little is known about the physiology and ecological significance of these associations. High concentrations of dimethylsulfoniopropionate (DMSP), a metabolite produced by marine microalgae, are commonly observed in coral-microalgae symbiosis, where DMS(P) is involved in multiple physiological functions. Knowledge on concentration and role of DMSP in analogous photosymbiosis in plankton is lacking. Here, we assess the total DMSP (DMSPt = DMSP + DMS) concentration and natural stable isotopes of sulfur across ecologically relevant symbiotic plankton groups, the Radiolaria and Foraminifera. We found that intracellular DMSPt concentrations in microalgal symbionts were among the highest recorded (range = 170–702 mmol L−1), while lower concentrations (range = 0.1–23 mmol L−1) were characteristic of the holobiont (i.e., host-microalgae). The contribution of symbiotic Radiolaria to the water column particulate DMSPt concentration ranged 0.1–8%. Sulfur isotopic composition (34S) of DMSPt in the Collodaria holobionts was significantly higher than their symbiotic microalgae isolated in culture. Despite their high intracellular DMSPt content, math formula uptake in these holobionts throughout 3-d incubations was not detected. We observed a systematic 34S depletion (∼ 1.5‰) of DMS relative to DMSP in experimental incubations containing filtered seawater, which we hypothesize is related to the bacterial preference for the uptake of 34S-depleted DMS. Overall, the results indicate that plankton symbiosis can, at times, represent a potentially important source of DMS(P). Specific differences in 34S provided new insights into sulfur isotopic fractionation associated with DMS(P) biotransformation processes, with potential implications for current interpretations of isotopically tracked biogenic sources of marine aerosols.

Lake Baikal, Siberia, is the most biodiverse freshwater lake on Earth. However, despite decades of painstaking limnological research on Baikal, broad spatial data on nutrient (nitrogen (N), phosphorus (P), silica (Si)) concentrations and temperature are sparse, as is our understanding of the bottom-up factors that limit phytoplankton in the lake. Earlier studies have suggested both N and P as limiting nutrients in Baikal, but the evidence, mostly based on elemental ratios, is limited and somewhat conflicting. We present experimental evidence that N and P co-limit phytoplankton productivity in some areas of Baikal during summer, along with the results of a comprehensive spatial survey of surface temperature, nutrients and chlorophyll a (Chl a) in Lake Baikal that support the experimental finding of colimitation. Surface water incubations from two trophically contrasting locations revealed co-limitation by N and P, as well as a positive effect of temperature (fluorescence after 5 d was ∼10% higher at 15°C than at 10°C). In a linear model of the survey data (26 sampling locations), N, P, and their interaction (N × P) were all significant predictors of Chl a concentration, indicating that either N or P (or both) may limit summer phytoplankton, depending on location. In contrast to the incubation experiments, temperature was not a significant predictor of Chl a concentration across the 26 sites we sampled. Lake Baikal is undergoing rapid warming and increased nutrient loading, which may boost phytoplankton productivity in the lake; however, the magnitude of this response will depend on ratios of soluble N and P inputs.

The foundation of methylmercury (MeHg) bioaccumulation in aquatic systems is uptake and trophic transfer into phytoplankton and zooplankton. Phytoplankton and zooplankton samples were collected seasonally from Long Island Sound (LIS) and its adjacent shelf waters, located along the northeast coast of the United States. Phytoplankton samples were size fractioned into 0.2–5 μm, 5–20 μm, and seston of > 20 μm. Average phytoplankton MeHg concentrations were 0.01–1.5 pmol g−1 (wet weight), and MeHg bioconcentration factors (logBCF) ranged from 2.5 to 5.5. There was higher MeHg concentration and logBCF values with increasing phytoplankton size. Zooplankton samples were size fractioned into 0.2–0.5 mm, 0.5–1.0 mm, 1.0–2.0 mm, and > 2.0 mm classes and analyzed for Hg and MeHg as well as carbon (δ13C), nitrogen (δ15N), and sulfur (δ34S) isotopes. The %MeHg in organisms was highest in the > 2.0 mm size class for coastal LIS and shelf species, indicating higher MeHg bioaccumulation with increasing zooplankton sizes. Eastern LIS zooplankton populations yielded higher overall Hg accumulation than western LIS populations, in alignment with phytoplankton fractions. Seasonal differences were evident for plankton %MeHg, although only ELIS displayed an increase in summertime %MeHg throughout the fractions. The results suggest the importance of biodilution in determining plankton MeHg concentrations. The δ13C values infer that zooplankton were opportunistically feeding on food containing variable levels of Hg and MeHg. The δ15N isotopes indicated enrichment characteristic of anthropogenic input in populations at the western end of LIS. The δ34S isotopes signified that organisms were primarily feeding on pelagic plankton.