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Ciliates have been recognized as one of the major components of the microbial food web, especially in ultra-oligotrophic waters, such as the Eastern Mediterranean Sea, where nutrients are scarce and the microbial community is dominated by pico- and nano-sized organisms. For this reason, ciliates play an important role in these ecosystems since they are the main planktonic grazers. Regardless the importance of these organisms, little is known about the community structure of heterotrophic and mixotrophic ciliates and how they are associated to their potential prey. In this study, we used 18S V4 rRNA gene metabarcoding to analyze ciliate community dynamics and how the relationship with potential prey changes according to different seasons and depths. Samples were collected seasonally at two stations of the Eastern Mediterranean Sea (HCB: coastal, M3A: offshore) from the surface and deep chlorophyll maximum (DCM) layers. The ciliate community structure varied across depths in HCB and across seasons in M3A, and the network analysis showed that in both stations, mixotrophic oligotrichs were positively associated with diatoms and showed few negative associations with ASVs annotated as marine Stramenopiles (MAST). On the other hand, heterotrophic tintinnids showed negative relationships in both HCB and M3A stations, mostly with Ochrophyta and Chlorophyta. These results showed, in first place that, although the two stations are close to each other, the ciliate dynamics differed between them. Moreover, mixotrophic and heterotrophic ciliates may have different ecological niches since mixotrophic ciliates may be more selective compared to heterotrophic species regarding their prey. These findings are the first glimpse into an understanding of the dynamics between heterotrophic and mixotrophic ciliates and their role in microbial assemblages and dynamics of ultra-oligotrophic environments., Ciliates have been recognized as one of the major components of the microbial food web, especially in ultra-oligotrophic waters, such as the Eastern Mediterranean Sea, where nutrients are scarce and the microbial community is dominated by pico- and nano-sized organisms. For this reason, ciliates play an important role in these ecosystems since they are the main planktonic grazers. Regardless the importance of these organisms, little is known about the community structure of heterotrophic and mixotrophic ciliates and how they are associated to their potential prey. In this study, we used 18S V4 rRNA gene metabarcoding to analyze ciliate community dynamics and how the relationship with potential prey changes according to different seasons and depths. Samples were collected seasonally at two stations of the Eastern Mediterranean Sea (HCB: coastal, M3A: offshore) from the surface and deep chlorophyll maximum (DCM) layers. The ciliate community structure varied across depths in HCB and across seasons in M3A, and the network analysis showed that in both stations, mixotrophic oligotrichs were positively associated with diatoms and showed few negative associations with ASVs annotated as marine Stramenopiles (MAST). On the other hand, heterotrophic tintinnids showed negative relationships in both HCB and M3A stations, mostly with Ochrophyta and Chlorophyta. These results showed, in first place that, although the two stations are close to each other, the ciliate dynamics differed between them. Moreover, mixotrophic and heterotrophic ciliates may have different ecological niches since mixotrophic ciliates may be more selective compared to heterotrophic species regarding their prey. These findings are the first glimpse into an understanding of the dynamics between heterotrophic and mixotrophic ciliates and their role in microbial assemblages and dynamics of ultra-oligotrophic environments.

It remains unclear as to how mixoplankton (coupled phototrophy and phagotrophy in one cell) affects the estimation of grazing rates obtained from the widely used dilution grazing technique. To address this issue, we prepared laboratory-controlled dilution experiments with known mixtures of phyto-, protozoo-, and mixoplankton, operated under different light regimes and species combinations. Our results evidenced that chlorophyll is an inadequate proxy for phytoplankton when mixoplankton are present. Conversely, species-specific cellular counts could assist (although not fully solve) in the integration of mixoplanktonic activity in a dilution experiment. Moreover, cell counts can expose prey selectivity patterns and intraguild interactions among grazers. Our results also demonstrated that whole community approaches mimic reality better than single-species laboratory experiments. We also confirmed that light is required for protozoo- and mixoplankton to correctly express their feeding activity, and that overall diurnal grazing is higher than nocturnal. Thus, we recommend that a detailed examination of initial and final plankton communities should become routine in dilution experiments, and that incubations should preferably be started at the beginning of both day and night periods. Finally, we hypothesize that in silico approaches may help disentangle the contribution of mixoplankton to the community grazing of a given system

Ciliate conjugation is considered a rare event to encounter in the field and it is mostly reported from cultures. In this work, we describe a synchronized conjugation event of planktonic ciliates that was discovered twice; in September 2019, at two different locations in the Cretan Sea, Eastern Mediterranean, and in October 2020. In 2019, first, at 2 m depth of the coastal station POSEIDON-HCB, in samples fixed with acid Lugol and formaldehyde, we found 340 and 200 mating pairs L(-1)of different ciliate species, respectively; and second, at the Heraklion port, we found 220 mating pairs L-1 of Strombidinopsis sp. and 1960 mating pairs L-1 of Strombidium sp. At the Heraklion port visited again in 2020, we found 800 mating pairs L-1 of Strombidinopsis sp. and 200 mating pairs L-1 of Strombidium sp. Since detailed descriptions of conjugation in pelagic oligotrich ciliates are missing, our observations indicate that ciliate conjugation could be a frequent and periodic phenomenon, under specific conditions.

info:eu-repo/semantics/published

Pollution of the marine environment is an emerging threat. Nowadays, engineered nanoparticles (<100 nm) such as zinc, copper and silver are widely used as antimicrobial agents, therefore often present in daily-life products. Consequently, the demand and production of nanoparticles are expected to increase. Here, we specifically focus on silver nanoparticles (AgNP). Once released into the environment, AgNPs pose an obvious ecotoxicological risk, potentially affecting ecosystem structure and functioning. For instance, phytoplankton-derived exudates, rich in acidic polysaccharides and amino acids, can abiotically aggregate into microgels such as transparent exopolymer particles (TEP) and Coomassie stainable particles (CSP). Hence, microgels can bridge dissolved and particulate size fractions and facilitate aggregate formation with organic and mineral particles. Both physical and chemical properties make TEP and CSP attractive nutrient hotspots for heterotrophic bacterioplankton. Bacteria, in turn, utilize extracellular enzymes to access these carbon and nitrogen pools. However, knowledge about the mechanisms by which AgNPs might interact with and affect the biogeochemical cycling of TEP and CSP is still insufficient. Therefore, we conducted a mesocosm experiment in the Eastern Mediterranean Sea and investigated the effects of environmentally relevant concentrations of silver ions (Ag+) and AgNP on the properties of TEP and CSP (i.e., area and abundance) along with enzymatic activity measurements. Our results showed that cyanobacteria were likely the primary source of CSP in the ultra-oligotrophic Mediterranean Sea. Also, CSP contributed more to the microgel pool than TEP, as indicated by a strong relationship between CSP and heterotrophic microbial dynamics. While silver (i.e., Ag+ or AgNP) had overall only marginal effects, both species affected the relationships between cell-specific LAPase activity and CSP and cell-specific APase activity and phosphate levels. Thus

The temporal variability of planktonic ciliates was studied in an extreme oligotrophic environment with special focus on trophic modes and size classes. Abundance, biomass, size classes, mixotrophy vs. heterotrophy, and species composition of planktonic ciliates were investigated focusing on temporal (samples collected on a monthly basis during 2019) and vertical (7 depth layers in the euphotic zone, from surface to 120 m) distribution at a coastal station in the oligotrophic Eastern Mediterranean. Abundance was in general very low (20 to 1150 cells L–1), except for September, which presented the highest abundance and biomass. Aloricate species dominated the ciliate community in all months and depths (% contribution from 77% in September to 99% in April). In general, oligotrichs presented maximum abundance at 2–10 m (except for June, July, and November: 100–120 m) whereas choreotrichs were more homogeneously distributed [and showed maxima at deep chlorophyll maximum (DCM)]. Small heterotrophs dominated the ciliate community at all depths and months, on average by 76% (they were 3 times more abundant than mixotrophs in terms of abundance and 2.5 times in terms of biomass). They were equally distributed both vertically and seasonally (and also in terms of size classes). In contrast, mixotrophs were found mainly at the surface layer to 20 m throughout the year, except for June and July (max at 100, 50 m). On average, 63% of integrated aloricate abundance was species <30 μm, of which 25% were mixotrophs. During the stratification period of May to November, the very small (<18 μm) and small (18–30 μm) mixotrophic species were distributed throughout the water column whereas the >50 μm mixotrophic species were found only above the DCM. In contrast, during the mixing period of December to May, mixotrophic ciliates were very few and were dominated by small and medium-sized species. It seems that mixotrophic and heterotrophic ciliates, on one hand, and

It remains unclear as to how mixoplankton (coupled phototrophy and phagotrophy in one cell) affects the estimation of grazing rates obtained from the widely used dilution grazing technique. To address this issue, we prepared laboratory-controlled dilution experiments with known mixtures of phyto-, protozoo-, and mixoplankton, operated under different light regimes and species combinations. Our results evidenced that chlorophyll is an inadequate proxy for phytoplankton when mixoplankton are present. Conversely, species-specific cellular counts could assist (although not fully solve) in the integration of mixoplanktonic activity in a dilution experiment. Moreover, cell counts can expose prey selectivity patterns and intraguild interactions among grazers. Our results also demonstrated that whole community approaches mimic reality better than single-species laboratory experiments. We also confirmed that light is required for protozoo- and mixoplankton to correctly express their feeding activity, and that overall diurnal grazing is higher than nocturnal. Thus, we recommend that a detailed examination of initial and final plankton communities should become routine in dilution experiments, and that incubations should preferably be started at the beginning of both day and night periods. Finally, we hypothesize that in silico approaches may help disentangle the contribution of mixoplankton to the community grazing of a given system

It remains unclear as to how mixoplankton (coupled phototrophy and phagotrophy in one cell) affects the estimation of grazing rates obtained from the widely used dilution grazing technique. To address this issue, we prepared laboratory-controlled dilution experiments with known mixtures of phyto-, protozoo-, and mixoplankton, operated under different light regimes and species combinations. Our results evidenced that chlorophyll is an inadequate proxy for phytoplankton when mixoplankton are present. Conversely, species-specific cellular counts could assist (although not fully solve) in the integration of mixoplanktonic activity in a dilution experiment. Moreover, cell counts can expose prey selectivity patterns and intraguild interactions among grazers. Our results also demonstrated that whole community approaches mimic reality better than single-species laboratory experiments. We also confirmed that light is required for protozoo- and mixoplankton to correctly express their feeding activity, and that overall diurnal grazing is higher than nocturnal. Thus, we recommend that a detailed examination of initial and final plankton communities should become routine in dilution experiments, and that incubations should preferably be started at the beginning of both day and night periods. Finally, we hypothesize that in silico approaches may help disentangle the contribution of mixoplankton to the community grazing of a given system.