Mixotrophic Protists Research Articles

Thermal response of freshwater ciliates: Can they survive at elevated lake temperatures?

Abstract Ciliates are a well‐defined, species‐rich, versatile group of heterotrophic and mixotrophic protists. Their response to increased temperature during global warming is critical for the structure and functioning of freshwater food webs. I conducted a meta‐analysis of the literature from field studies and experimental evidence to assess the parameters characterising the thermal response of freshwater ciliates. The shape of the thermal performance curve predicts the ciliates' survival at supraoptimal temperatures (i.e., the width of the thermal safety margin). The ciliates' typical thermal safety margin is c. 5°C. More than two‐thirds of the freshwater ciliates dwelling permanently or occasionally in the pelagial cannot survive at temperatures exceeding 30°C. Likewise, cold‐stenothermic species, which represent a significant fraction of euplanktonic ciliates, cannot survive by evolutionary adaptation to rapidly warming environments. The statistical analysis revealed that the ciliates' thermal performance is affected by their planktonic lifestyle (euplanktonic vs. tychoplanktonic), ability to form cysts and nutritional ecology. Bactivorous ciliates have the widest temperature niche, and algivorous ciliates have the narrowest temperature niche. Phenotypic plasticity and genetic variation, favouring the selection of pre‐adapted species in a new environment, are widespread among freshwater ciliates. However, the lack of evidence for the temperature optima and imprecisely defined tolerance limits of most species hampers the present analysis. The extent of acclimation and adaptation requires further research with more ciliate species than the few chosen thus far. Recent eco‐evolutionary experimental work and modelling approaches have demonstrated that the ciliates' thermal responses follow general trends predicted by the metabolic theory of ecology and mechanistic functions inherent in enzyme kinetics. The present analysis identified current knowledge gaps and avenues for future research that may serve as a model study for other biota. Thermal adaptation may conflict with adaptation to other stressors (predators, food availability, pH), making general predictions on the future role of freshwater ciliates in a warmer environment difficult, if not impossible, at present.

Integrating depth-dependent protist dynamics and microbial interactions in spring succession of a freshwater reservoir

BackgroundProtists are essential contributors to eukaryotic diversity and exert profound influence on carbon fluxes and energy transfer in freshwaters. Despite their significance, there is a notable gap in research on protistan dynamics, particularly in the deeper strata of temperate lakes. This study aimed to address this gap by integrating protists into the well-described spring dynamics of Římov reservoir, Czech Republic. Over a 2-month period covering transition from mixing to established stratification, we collected water samples from three reservoir depths (0.5, 10 and 30 m) with a frequency of up to three times per week. Microbial eukaryotic and prokaryotic communities were analysed using SSU rRNA gene amplicon sequencing and dominant protistan groups were enumerated by Catalysed Reporter Deposition-Fluorescence in situ Hybridization (CARD-FISH). Additionally, we collected samples for water chemistry, phyto- and zooplankton composition analyses.ResultsFollowing the rapid changes in environmental and biotic parameters during spring, protistan and bacterial communities displayed swift transitions from a homogeneous community to distinct strata-specific communities. A prevalence of auto- and mixotrophic protists dominated by cryptophytes was associated with spring algal bloom-specialized bacteria in the epilimnion. In contrast, the meta- and hypolimnion showcased a development of a protist community dominated by putative parasitic Perkinsozoa, detritus or particle-associated ciliates, cercozoans, telonemids and excavate protists (Kinetoplastida), co-occurring with bacteria associated with lake snow.ConclusionsOur high-resolution sampling matching the typical doubling time of microbes along with the combined microscopic and molecular approach and inclusion of all main components of the microbial food web allowed us to unveil depth-specific populations’ successions and interactions in a deep lentic ecosystem.

Contrasting responses of different mixotrophic protists to light and nutrient availability

AbstractMixotrophic protists are important members of aquatic microbial food webs where they can dominate bacterivory and strongly impact energy and nutrient flow. While light and nutrient availability are known to impact grazing rates by mixotrophs in laboratory studies, little is known about how changes in resource availability affect mixotrophic organisms in natural communities. Here, we performed a short‐term grazing assay using fluorescently labeled bacteria in a phosphorus‐limited subalpine mountain lake to test how the availability of light and nutrients affect mixotroph grazing rates. High grazing rates by a Pseudopedinella‐like dictyochophyte were unaffected by nutrient or light manipulation. In contrast, grazing rates by pigmented cryptophytes significantly decreased upon phosphorus addition, light deprivation, and a combination of both. We compare these findings with the seasonal analysis of a 4‐yr time‐series dataset of 18S rRNA gene amplicon abundances. While cryptophytes dominated in autumn and winter and were negatively related with water column stability, Pseudopedinella amplicon abundances showed a consistent spring–summer peak and were positively related with chlorophyll a concentrations. These results demonstrate the diversity in nutritional strategies among co‐occurring mixotrophic protists, which cannot be resolved by quantification of bulk grazing rates. Moreover, these different strategies result in distinct seasonal dynamics and coexistence of different mixotrophs, suggesting their respective trajectories into future aquatic ecosystems will also differ.

Environment-dependent metabolic investments in the mixotrophic chrysophyte Ochromonas.

Mixotrophic protists combine photosynthesis and phagotrophy to obtain energy and nutrients. Because mixotrophs can act as either primary producers or consumers, they have a complex role in marine food webs and biogeochemical cycles. Many mixotrophs are also phenotypically plastic and can adjust their metabolic investments in response to resource availability. Thus, a single species's ecological role may vary with environmental conditions. Here, we quantified how light and food availability impacted the growth rates, energy acquisition rates, and metabolic investment strategies of eight strains of the mixotrophic chrysophyte, Ochromonas. All eight Ochromonas strains photoacclimated by decreasing chlorophyll content as light intensity increased. Some strains were obligate phototrophs that required light for growth, while other strains showed stronger metabolic responses to prey availability. When prey availability was high, all eight strains exhibited accelerated growth rates and decreased their investments in both photosynthesis and phagotrophy. Photosynthesis and phagotrophy generally produced additive benefits: In low-prey environments, Ochromonas growth rates increased to maximum, light-saturated rates with increasing light but increased further with the addition of abundant bacterial prey. The additive benefits observed between photosynthesis and phagotrophy in Ochromonas suggest that the two metabolic modes provide nonsubstitutable resources, which may explain why a tradeoff between phagotrophic and phototrophic investments emerged in some but not all strains.

Implementation and assessment of a model including mixotrophs and the carbonate cycle (Eco3M_MIX-CarbOx v1.0) in a highly dynamic Mediterranean coastal environment (Bay of Marseille, France) – Part 1: Evolution of ecosystem composition under limited light and nutrient conditions

Abstract. Many current biogeochemical models rely on an autotrophic versus heterotrophic food web representation. However, in recent years, an increasing number of studies have begun to challenge this approach. Several authors have highlighted the importance of protists capable of combining photoautotrophic and heterotrophic nutrition in a single cell. These mixotrophic protists are known to play an important role in the carbon cycle. Here, we present a new biogeochemical model that represents the food web using variable stoichiometry. It contains the classic compartments such as zooplankton, phytoplankton, and heterotrophic bacteria and a newly added compartment to represent two types of mixotrophic protists: non-constitutive mixotrophs (NCMs) and constitutive mixotrophs (CMs). We demonstrate that the model correctly reproduces the characteristics of NCMs and CMs and proceed to study the impact of light and nutrient limitation on planktonic ecosystem structure in a highly dynamic Mediterranean coastal area, namely the Bay of Marseille (BoM, France), paying special attention to the dynamics of mixotrophic protists in these limiting conditions. In addition, we investigate the carbon, nitrogen, and phosphorus fluxes associated with mixotrophic protists and showed the following: (i) the portion of the ecosystem in terms of the percentage of carbon biomass occupied by NCMs decreases when resources (nutrient and prey concentrations) decrease, although their mixotrophy allows them to maintain a carbon biomass almost as significant as the copepod one (129.8 and 148.7 mmol C m−3, respectively), as photosynthesis increases as a food source, and (ii) the portion of the ecosystem in terms of the percentage of carbon biomass occupied by CM increases when nutrient concentrations decrease due to their capability to ingest prey to supplement their N and P needs. In addition to providing new insights regarding the conditions that lead to the emergence of mixotrophs in the BoM, this work provides a new tool to perform long-term studies and predictions of mixotroph dynamics in coastal environments under different environmental forcings.

Disentangling the role of light and nutrient limitation on bacterivory by mixotrophic nanoflagellates.

Many phytoplankton taxa function on multiple trophic levels by combining photosynthesis and ingestion of bacteria, termed mixotrophy. Despite the recognition of mixotrophy as a universal functional trait, we have yet to fully resolve how environmental conditions influence community grazing rates in situ. A microcosm study was used to assess bacterivory by mixotrophic nanoflagellates following nutrient enrichment and light attenuation in a temperate lake. We found contrasting results based on assessment of mixotroph abundance or bacterivory. Despite an interactive effect of nutrient enrichment and light attenuation on mixotroph abundance, significant differences within light treatments were observed only after enrichment with P or N + P. The greatest abundance of mixotrophs across treatments occurred under co-nutrient enrichment with full exposure to irradiance. However, bacterivory by mixotrophic nanoflagellates was greatest under shaded conditions after either N or P enrichment. We suggest that PAR availability dampened the stimulatory effect of nutrient limitation, and bacterivory supplemented a suboptimal photosynthetic environment. In a saturating light regime, the mixotrophic community was less driven to ingest bacteria because photosynthesis was able to satisfy energetic demands. These findings quantify community bacterivory in response to environmental drivers that may characterize future ecosystem conditions and highlight the importance of considering grazing rates in conjunction with abundance of mixotrophic protists.

Winners and Losers of Atlantification: The Degree of Ocean Warming Affects the Structure of Arctic Microbial Communities.

Arctic microbial communities (i.e., protists and bacteria) are increasingly subjected to an intrusion of new species via Atlantification and an uncertain degree of ocean warming. As species differ in adaptive traits, these oceanic conditions may lead to compositional changes with functional implications for the ecosystem. In June 2021, we incubated water from the western Fram Strait at three temperatures (2 °C, 6 °C, and 9 °C), mimicking the current and potential future properties of the Arctic Ocean. Our results show that increasing the temperature to 6 °C only minorly affects the community, while an increase to 9 °C significantly lowers the diversity and shifts the composition. A higher relative abundance of large hetero- and mixotrophic protists was observed at 2 °C and 6 °C compared to a higher abundance of intermediate-sized temperate diatoms at 9 °C. The compositional differences at 9 °C led to a higher chlorophyll a:POC ratio, but the C:N ratio remained similar. Our results contradict the common assumption that smaller organisms and heterotrophs are favored under warming and strongly indicate a thermal limit between 6 °C and 9 °C for many Arctic species. Consequently, the magnitude of temperature increase is a crucial factor for microbial community reorganization and the ensuing ecological consequences in the future Arctic Ocean.

Trophic strategies explain the ocean niches of small eukaryotic phytoplankton.

A large fraction of marine primary production is performed by diverse small protists, and many of these phytoplankton are phagotrophic mixotrophs that vary widely in their capacity to consume bacterial prey. Prior analyses suggest that mixotrophic protists as a group vary in importance across ocean environments, but the mechanisms leading to broad functional diversity among mixotrophs, and the biogeochemical consequences of this, are less clear. Here we use isolates from seven major taxa to demonstrate a tradeoff between phototrophic performance (growth in the absence of prey) and phagotrophic performance (clearance rate when consuming Prochlorococcus). We then show that trophic strategy along the autotrophy-mixotrophy spectrum correlates strongly with global niche differences, across depths and across gradients of stratification and chlorophyll a. A model of competition shows that community shifts can be explained by greater fitness of faster-grazing mixotrophs when nutrients are scarce and light is plentiful. Our results illustrate how basic physiological constraints and principles of resource competition can organize complexity in the surface ocean ecosystem.

Taxonomic Composition of Protist Communities in the Coastal Stratified Lake Kislo-Sladkoe (Kandalaksha Bay, White Sea) Revealed by Microscopy

Lake Kislo-Sladkoe is a stratified water body partly isolated from the White Sea. Perennial meromixis in the lake irregularly alternates with mixing events. Taking into account that the protists of Arctic coastal stratified water bodies are understudied, we evaluated for the first time the vertical structure, species richness, and diversity of protists assigned to different taxonomic groups in Lake Kislo-Sladkoe using light, luminescent, and scanning electron microscopy. To test the research hypothesis that a mixing event affects the vertical stratification and species composition of protists in a stratified lake, we compared the protist communities of Lake Kislo-Sladkoe in two extremely different states: strong meromixis vs. full vertical mixing. A total of 97 morphologically distinct phototrophic, heterotrophic, and mixotrophic protists were revealed with the most diverse supertaxa SAR (59), Obazoa (9), and Excavates (14). The hidden diversity of protists (43 species) was a bit less than the active diversity (54 species). A taxonomic list and micrographs of cells for the observed protists are provided. The majority of species revealed are cosmopolitan or widespread in the northern sea waters. The vertical patterns of protist communities were absolutely different in 2018 and 2021. In July 2018, clearly distinct protist communities inhabited different layers of the lake. Bloom of cryptophyte Rhodomonas cf. baltica was detected in chemocline, whereas the maximum density of its grazers was observed in adjacent layers, mainly dinoflagellates Gymnodinium sp. and Scrippsiella trochoidea, as well as a ciliate Prorodon sp. In 2021 due to the recent mixing of lake and seawater, there were no distinct communities in the water column except the superficial 0–1 m layer of fresh water.

Apparent shifts in the microplankton community in response to zinc and iron enrichments in the Northeastern Subarctic Pacific

The offshore Gulf of Alaska—in the northeastern subarctic Pacific—is a high-nitrate, low-chlorophyll (HNLC) oceanic region where concentrations of dissolved iron (DFe) and zinc (DZn) are of the order ~0.1 nmol kg−1 in surface waters. At the average winter mixed layer depth of ~120 m, DFe and DZn are of the orders ~0.1–0.2 nmol kg−1 and ~1–2 nmol kg−1, respectively. Vertical supply of Fe to surface waters is therefore limited, with phytoplankton blooms driven by episodic atmospheric and lateral inputs of Fe. In contrast, DZn is presumably replenished through seasonal vertical mixing, though maintained at low levels in surface waters, even in winter, implying that removal processes are in operation. Previous supplementation experiments have demonstrated that phytoplankton biomass (chl-a), growth, and drawdown of nitrate (NO3-) are stimulated strongly by Fe but only minimally, if at all, by Zn alone. Using on-deck bioassay incubations of surface waters, we confirm that the addition of Fe stimulates significant increases in chl-a and large diatoms, and drawdown of NO3- and silicic acid [Si(OH)4]. Associated drawdown of DZn and dissolved cadmium (DCd) indicated Zn stress in the control and +Fe treatment. Supplementation with Zn alone had no significant impact on NO3- and commonly monitored pigments such as chl-a and fucoxanthin. However, in the +Zn treatment, DFe and coccolithophore abundance were significantly lower, and the concentration of particulate organic carbon (POC) and that of the pigments alloxanthin and chlorophyll c1+2 were significantly higher than those in control incubations. Our experiment corroborates previously observed relationships between DZn and alloxanthin and chlorophyll c across the subarctic north Pacific and Bering Sea and could help to explain the low or undetectable concentrations of alloxanthin frequently observed in this region. Alloxanthin is a pigment specific to the cryptophytes, either within free-living cells or within functional plastids retained by mixotrophic protists. Both cryptophytes and mixotrophic ciliates, such as Laboea sp. and Strombidium spp., make a variable but significant contribution to phytoplankton biomass in this area. In the absence of Fe supply to surface waters, Zn could play a hitherto unsuspected role in shaping plankton communities in the HNLC Gulf of Alaska.

Light dependence in the phototrophy-phagotrophy balance of constitutive and non-constitutive mixotrophic protists.

Mixotrophic protists display contrasting nutritional strategies and are key groups connecting planktonic food webs. They comprise constitutive mixotrophs (CMs) that have an innate photosynthetic ability and non-constitutive mixotrophs (NCMs) that acquire it from their prey. We modelled phototrophy and phagotrophy of two mixotrophic protists as a function of irradiance and prey abundance. We hypothesised that differences in their physiology (constitutive versus non-constitutive mixotrophy) can result in different responses to light gradients. We fitted the models with primary production and bacterivory data from laboratory and field experiments with the nanoflagellate Chrysochromulina parva (CM) and the ciliate Ophrydium naumanni (NCM) from north Andean Patagonian lakes. We found a non-monotonic response of phototrophy and phagotrophy to irradiance in both mixotrophs, which was successfully represented by our models. Maximum values for phototrophy and phagotrophy were found at intermediate irradiance coinciding with the light at the deep chlorophyll maxima in these lakes. At lower and higher irradiances, we found a decoupling between phototrophy and phagotrophy in the NCM while these functions were more coupled in the CM. Our modelling approach revealed the difference between both mixotrophic functional types on the balance between their nutritional strategies under different light scenarios. Thus, our proposed models can be applied to account how changing environmental conditions affect both primary and secondary production within the planktonic microbial food web.

Mixotrophy in the bloom-forming genus Phaeocystis and other haptophytes

Phaeocystis is a globally widespread marine phytoplankton genus, best known for its colony-forming species that can form large blooms and odorous foam during bloom decline. In the North Sea, Phaeocystis globosa typically becomes abundant towards the end of the spring bloom, when nutrients are depleted and the share of mixotrophic protists increases. Although mixotrophy is widespread across the eukaryotic tree of life and is also found amongst haptophytes, a mixotrophic nutrition has not yet been demonstrated in Phaeocystis. Here, we sampled two consecutive Phaeocystis globosa spring blooms in the coastal North Sea. In both years, bacterial cells were observed inside 0.6 – 2% of P. globosa cells using double CARD-FISH hybridizations in combination with laser scanning confocal microscopy. Incubation experiments manipulating light and nutrient availability showed a trend towards higher occurrence of intracellular bacteria under P-deplete conditions. Based on counts of bacteria inside P. globosa cells in combination with theoretical values of prey digestion times, maximum ingestion rates of up to 0.08 bacteria cell−1 h−1 were estimated. In addition, a gene-based predictive model was applied to the transcriptome assemblies of seven Phaeocystis strains and 24 other haptophytes to assess their trophic mode. This model predicted a phago-mixotrophic feeding strategy in several (but not all) strains of P. globosa, P. antarctica and other haptophytes that were previously assumed to be autotrophic. The observation of bacterial cells inside P. globosa and the gene-based model predictions strongly suggest that the phago-mixotrophic feeding strategy is widespread among members of the Phaeocystis genus and other haptophytes, and might contribute to their remarkable success to form nuisance blooms under nutrient-limiting conditions.

Complex Trophic Interactions in an Acidophilic Microbial Community.

Extreme habitats often harbor specific communities that differ substantially from non-extreme habitats. In many cases, these communities are characterized by archaea, bacteria and protists, whereas the number of species of metazoa and higher plants is relatively low. In extremely acidic habitats, mostly prokaryotes and protists thrive, and only very few metazoa thrive, for example, rotifers. Since many studies have investigated the physiology and ecology of individual species, there is still a gap in research on direct, trophic interactions among extremophiles. To fill this gap, we experimentally studied the trophic interactions between a predatory protist (Actinophrys sol, Heliozoa) and its prey, the rotifers Elosa woralli and Cephalodella sp., the ciliate Urosomoida sp. and the mixotrophic protist Chlamydomonas acidophila (a green phytoflagellate, Chlorophyta). We found substantial predation pressure on all animal prey. High densities of Chlamydomonas acidophila reduced the predation impact on the rotifers by interfering with the feeding behaviour of A. sol. These trophic relations represent a natural case of intraguild predation, with Chlamydomonas acidophila being the common prey and the rotifers/ciliate and A. sol being the intraguild prey and predator, respectively. We further studied this intraguild predation along a resource gradient using Cephalodella sp. as the intraguild prey. The interactions among the three species led to an increase in relative rotifer abundance with increasing resource (Chlamydomonas) densities. By applying a series of laboratory experiments, we revealed the complexity of trophic interactions within a natural extremophilic community.

Selective Feeding of a Mixotrophic Dinoflagellate (Lepidodinium sp.) in Response to Experimental Warming and Inorganic Nutrient Imbalance.

Mixotrophic protists are widely observed in the aquatic ecosystems, while how they respond to inorganic nutrient imbalance and ocean warming remains understudied. We conducted a series of experiments on a mixotrophic dinoflagellate Lepidodinium sp. isolated from subtropical coastal waters to investigate the combined effect of temperature and medium nitrate to phosphate ratio (N:P ratio) on the ingestion activities of mixotrophic protists. We found Lepidodinium sp. displayed selective feeding behaviour with a higher ingestion rate on high-N prey (N-rich Rhodomonas salina) when the ambient inorganic N:P ratio was equal to or below the Redfield ratio. The Chesson selectivity index α increased with increasing temperature, suggesting that warming exacerbated the selective feeding of Lepidodinium sp. Under inorganic nitrogen sufficient conditions (N:P ratio = 64), no selective feeding was observed at 25 and 28°C, while it occurs at 31°C, which also indicates that warming alters the feeding behaviour of Lepidodinium sp. In addition, our results revealed that the total ingestion rate of Lepidodinium sp. under the condition with normal inorganic nutrients (Redfield ratio) was significantly lower than that under nutrient-imbalanced conditions, which indicates that Lepidodinium sp. developed compensatory feeding to balance their cellular stoichiometry and satisfy their growth. Our study is the first attempt on revealing the selective feeding behaviours of mixotrophic protists on prey under different inorganic nutrient environments and rising temperatures, which will contribute to our understanding of the response of marine plankton food web to projected climate changes.

Mucospheres produced by a mixotrophic protist impact ocean carbon cycling

Mixotrophic protists (unicellular eukaryotes) that engage in both phototrophy (photosynthesis) and phago-heterotrophy (engulfment of particles)—are predicted to contribute substantially to energy fluxes and marine biogeochemical cycles. However, their impact remains largely unquantified. Here we describe the sophisticated foraging strategy of a widespread mixotrophic dinoflagellate, involving the production of carbon-rich ‘mucospheres’ that attract, capture, and immobilise microbial prey facilitating their consumption. We provide a detailed characterisation of this previously undescribed behaviour and reveal that it represents an overlooked, yet quantitatively significant mechanism for oceanic carbon fluxes. Following feeding, the mucospheres laden with surplus prey are discarded and sink, contributing an estimated 0.17–1.24 mg m−2 d−1 of particulate organic carbon, or 0.02–0.15 Gt to the biological pump annually, which represents 0.1–0.7% of the estimated total export from the euphotic zone. These findings demonstrate how the complex foraging behaviour of a single species of mixotrophic protist can disproportionally contribute to the vertical flux of carbon in the ocean.

The dynamic trophic architecture of open-ocean protist communities revealed through machine-guided metatranscriptomics

Intricate networks of single-celled eukaryotes (protists) dominate carbon flow in the ocean. Their growth, demise, and interactions with other microorganisms drive the fluxes of biogeochemical elements through marine ecosystems. Mixotrophic protists are capable of both photosynthesis and ingestion of prey and are dominant components of open-ocean planktonic communities. Yet the role of mixotrophs in elemental cycling is obscured by their capacity to act as primary producers or heterotrophic consumers depending on factors that remain largely uncharacterized. Here, we develop and apply a machine learning model that predicts the in situ trophic mode of aquatic protists based on their patterns of gene expression. This approach leverages a public collection of protist transcriptomes as a training set to identify a subset of gene families whose transcriptional profiles predict trophic mode. We applied our model to nearly 100 metatranscriptomes obtained during two oceanographic cruises in the North Pacific and found community-level and population-specific evidence that abundant open-ocean mixotrophic populations shift their predominant mode of nutrient and carbon acquisition in response to natural gradients in nutrient supply and sea surface temperature. Metatranscriptomic data from ship-board incubation experiments revealed that abundant mixotrophic prymnesiophytes from the oligotrophic North Pacific subtropical gyre rapidly remodeled their transcriptome to enhance photosynthesis when supplied with limiting nutrients. Coupling this approach with experiments designed to reveal the mechanisms driving mixotroph physiology provides an avenue toward understanding the ecology of mixotrophy in the natural environment.

Housekeeping in the Hydrosphere: Microbial Cooking, Cleaning, and Control under Stress.

Who’s cooking, who’s cleaning, and who’s got the remote control within the waters blanketing Earth? Anatomically tiny, numerically dominant microbes are the crucial “homemakers” of the watery household. Phytoplankton’s culinary abilities enable them to create food by absorbing sunlight to fix carbon and release oxygen, making microbial autotrophs top-chefs in the aquatic kitchen. However, they are not the only bioengineers that balance this complex household. Ubiquitous heterotrophic microbes including prokaryotic bacteria and archaea (both “bacteria” henceforth), eukaryotic protists, and viruses, recycle organic matter and make inorganic nutrients available to primary producers. Grazing protists compete with viruses for bacterial biomass, whereas mixotrophic protists produce new organic matter as well as consume microbial biomass. When viruses press remote-control buttons, by modifying host genomes or lysing them, the outcome can reverberate throughout the microbial community and beyond. Despite recognition of the vital role of microbes in biosphere housekeeping, impacts of anthropogenic stressors and climate change on their biodiversity, evolution, and ecological function remain poorly understood. How trillions of the smallest organisms in Earth’s largest ecosystem respond will be hugely consequential. By making the study of ecology personal, the “housekeeping” perspective can provide better insights into changing ecosystem structure and function at all scales.

Impact of nutrient availability on the trophic strategies of the planktonic protist communities in a disturbed Mediterranean coastal lagoon

The impact of changes in nitrogen (N) and phosphorus (P) availability on the trophic strategies of planktonic protists was evaluated in a disturbed Mediterranean lagoon (Biguglia lagoon, France) using short-term bioassays. Natural communities were collected in three periods, i.e., autumn, spring and summer, to address the influence of the different environmental conditions. The responses of autotrophic plankton communities to experimentally induced N and/or P limitations were assessed as changes in chlorophyll a (Chl a) concentrations and in the abundances of potentially mixotrophic protists taxa. We observed blooms (> 105 cells l−1) of nanoflagellates in autumn, and of phycocyanin-rich picocyanobacteria in summer. Communities showed a co-limitation by N and P at the three sampling periods, despite high N:P ratios in autumn and spring. The high abundances of potentially mixotrophic dinoflagellates during these periods suggest the involvement of alternative trophic pathways for their maintenance in the lagoon. After bioassay incubations using different nutrient enrichment treatments, we often observed reduced abundances of mixotrophic protists containing Chl a with a concomitant increased abundance of protists without Chl a. This indicates a loss of chloroplasts and photoautotrophic abilities in protists cells, possibly reflecting a shift towards heterotrophy that could be sustained by phagotrophy.

Growth and photosynthetic responses of Ochromonas gloeopara to cadmium stress and its capacity to remove cadmium.

Cadmium (Cd) is one of the predominant anthropogenic pollutants in aquatic systems. As Cd has negative effects on species at all trophic levels, the community composition in aquatic habitats can be changed as a result of Cd stress. The response of mixotrophic protists to environmental stressors is particularly important as they act as both producers and consumers in complex planktonic communities. In this study, we used mixotrophic Ochromonas gloeopara to study its growth and photosynthetic responses to Cd, and specially focused on the effects of initial Cd concentrations and nutrient levels on its capacity to remove Cd. Results showed that when Cd concentration reached 0.5mgL-1, the growth rate and carrying capacity were significantly inhibited, whereas the photosynthesis was markedly decreased when Cd concentration reached 0.15mgL-1. Moreover, under Cd concentration 0.15, 0.5, 0.9, 1.6, and 2.0mgL-1, the removal efficiencies of Cd by O.gloeopara were 83.2%, 77.7%, 74.6%, 70.1%, and 68.8%, respectively. The increase of nitrogen did not cause significant effect on the removal capacity of Cd by O.gloeopara, but increased concentration of phosphorus significantly enhanced the removal capacity of Cd. Our findings indicated that the mixotrophic O.gloeopara has strong tolerance and capacity to remove Cd, and increasing concentration of phosphorus can increase its removal capacity, suggesting that O.gloeopara has great potential application value in mitigating Cd pollution in waters.

Plasticity and Multiplicity of Trophic Modes in the Dinoflagellate Karlodinium and Their Pertinence to Population Maintenance and Bloom Dynamics

As the number of mixotrophic protists has been increasingly documented, “mixoplankton”, a third category separated from the traditional categorization of plankton into “phytoplankton” and “zooplankton”, has become a new paradigm and research hotspot in aquatic plankton ecology. While species of dinoflagellates are a dominant group among all recorded members of mixoplankton, the trophic modes of Karlodinium, a genus constituted of cosmopolitan toxic species, were reviewed due to their representative features as mixoplankton and harmful algal blooms (HABs)-causing dinoflagellates. Among at least 15 reported species in the genus, three have been intensively studied for their trophic modes, and all found to be phagotrophic. Their phagotrophy exhibits multiple characteristics: (1) omnivority, i.e., they can ingest a variety of preys in many forms; (2) flexibility in phagotrophic mechanisms, i.e., they can ingest small preys by direct engulfment and much bigger preys by myzocytosis using a peduncle; (3) cannibalism, i.e., species including at least K. veneficum can ingest the dead cells of their own species. However, for some recently described and barely studied species, their tropical modes still need to be investigated further regarding all of the above-mentioned aspects. Mixotrophy of Karlodinium plays a significant role in the population dynamics and the formation of HABs in many ways, which thus deserves further investigation in the aspects of physiological ecology, environmental triggers (e.g., levels of inorganic nutrients and/or presence of preys), energetics, molecular (genes and gene expression regulations) and biochemical (e.g., relevant enzymes and signal molecules) bases, origins, and evaluation of the advantages of being a phagotroph.