Ciliate Assemblages Research Articles

Two-Decade Changes in the Ciliate Assemblage Feeding Pattern Reflect the Reservoir Nutrient Load

The perception of the importance of ciliate in freshwater has changed dramatically since the “microbial loop” conceptualisation, reflecting methodological attempts. The data from two decades (1994–2018) on the surface (0–3 m) ciliate assemblage in the Slapy reservoir (Vltava River, Czech Republic) during two different nutrient-load defined periods were analysed. We grouped the identified, quantified, and biomass-evaluated ciliates in the quantitative protargol-impregnated preparations according to their feeding behaviour. The sampling median and interquartile range data of the ciliates were plotted; the modelled water age, nutrients, bacteria, heterotrophic nanoflagellates, and Rhodomonas spp. were applied as the main explanatory background variables. We validated the differences between the periods, engaging multivariate analyses. The picoplankton-filtering species dominated the assemblages in an annual mean (halteriids and minute strobilidiids followed by peritrichs). Algae hunting urotrichs, Balanion planctonicum, and nanoplankton filtering tintinnids were significant before the spring phytoplankton peak when a maximum of ciliate biomass reflected mixotrophic nanoplankton filtering pelagostrombidiids. Only there did ciliate biomass tightly follow their quantified prey. Heterotrophic and mixotrophic Askenasia and Lagynophrya were typical raptorial/flagellate-hunting cilates; only Mesodinium spp. reached the maximum during autumn. The observed oligotrophication of the reservoir increased the ciliate assemblage biomass in the surface layer during stratification in concordance with the Plankton Ecology Group (PEG) model.

Variability of winter cooling affects intensity of phytoplankton spring blooms – how resilient is the ciliate assemblage composition to changes in food availability?

After years of partial winter mixing in Lake Zurich (Switzerland), a complete turnover of the water column reoccurred during winter/spring 2021. It was favored by a cold, windy winter and a small difference of water temperatures between the surface zone and a hypolimnion (deep water zone) that had gradually warmed during the previous years. The trend of declining phytoplankton spring blooms due to incomplete winter mixing was interrupted by mass development of algae due to the upwelling of nutrients accumulated in the hypolimnion. The effects of this singular deep mixing on the microbial food web during spring were studied in a high-frequency sampling campaign and compared with data from two years of partial winter mixing (2020 and 2022). A particular focus was put on the quantitative composition of the ciliate assemblage. Our results showed that not all organisms reacted equally to the nutrient (phosphorus) boost in the surface zone. Centric diatoms and cryptophytes profited most directly from the deep mixing, outcompeting the otherwise dominant cyanobacterium Planktothrix rubescens. Heterotrophic bacteria and their top predators, the ‘heterotrophic nanoflagellates’ trophic guild, were less affected by the nutrient supply and showed only short-lived increases of maximal biomass. The assemblage composition of ciliate morphotypes was highly resilient over the three years, presumably due to the range of acceptable food items of the predominant omnivorous species. However, numerous ciliate morphotypes showed brief mass development in 2021, and Balanion planctonicum, small Urotricha species and tintinnids were significantly more frequent than in 2020/2022. Small interception-feeding morphotypes apparently profited from the rich supply of their cryptomonad food, and tintinnid morphotypes additionally benefited from the availability of building material (e.g., centric diatom shells) for their loricae. In summary, we show that effects of lake warming in deep stratifying lakes are not as unidirectional as previously presumed, and we reveal resilience of the pelagic ciliate morphotype assemblage to lake warming related interannual variability in Lake Zurich.

Connectivity of benthic marine ciliates assemblages in sandy beaches of south Bahia (Brazil)

Sandy beaches are predominant ecosystems in the coastal zone of Brazil. In the interstitial environment, several microscopic species called microbenthos are responsible for primary production, consumption of bacteria and decomposition processes. Ciliophora is one of the most diverse phylum within the microbenthos, although its diversity and ecological aspects are still underestimated. We selected 12 beach segments in the south coast of Bahia, northeastern Brazil, to verify the diversity and connectivity between open and extensive beaches and those isolated by rocky outcrops. We found 52 morphospecies distributed in 16 families of ciliates. Statistical analysis showed higher richness in open beaches when compared to isolated ones, that could be due to their extension area, low hydrodynamics and organic matter retention. Urbanization factor, when interacting with beach categories, affect ciliates species richness. In our samples, open beaches presented a higher number of ciliate families in comparison with isolated beaches, and urbanization seemed to influence these differences. Species composition analysis showed high connectivity of ciliate assemblages between the beaches sampled, without significant diferences. The diversity profile showed that open beaches have greater diversity in comparison with isolated beaches. Urbanization profiles did not show diferences in ciliate diversity. More studies on marine benthic ciliates diversity and their ecological aspects are needed to understand their distribution patterns and assemblage connectivity.

Influence of the antibiotic nitrofurazone on community dynamics of marine periphytic ciliates: Evidence from community-based bioassays

Marine periphytic ciliates play a pivotal role in shaping coastal ecosystems dynamics, thereby acting as robust biological indicators of aquatic ecosystem health and functionality. However, the understanding of the effects of veterinary antibiotics on composition and structure of periphytic ciliate communities remains limited. Therefore, this research investigates the influence of the veterinary antibiotic nitrofurazone on the community dynamics of marine periphytic ciliates through bioassay experiments conducted over a one-year cycle. Various concentrations of nitrofurazone were administered to the tested ciliate assemblages, and subsequent changes in community composition, abundance, and diversity were quantitatively analyzed. The research revealed significant alterations in periphytic ciliate communities following exposure to nitrofurazone. Concentration-dependent (0–8 mg L−1) decrease in ciliates abundance, accompanied by shifts in species composition, community structure, and community patterns were observed. Comprehensive assessment of diversity metrics indicated significant changes in species richness and evenness in the presence of nitrofurazone, potentially disrupting the stability of ciliate communities. Furthermore, nitrofurazone significantly influenced the community structure of ciliates in all seasons (winter: R2 = 0.489; spring: R2 = 0.666; summer: R2 = 0.700, autumn: R2 = 0.450), with high toxic potential in treatments 4 and 8 mg L−1. Differential abundances of ciliates varied across seasons and nitrofurazone treatments, some orders like Pleurostomatida were consistently affected, while others (i.e., Strombidida and Philasterida) showed irregular distributions or were evenly affected (e.g., Urostylida and Synhymeniida). Retrieved contrasting patterns between nitrofurazone and community responses underscore the broad response repertoire exhibited by ciliates to antibiotic exposure, suggesting potential cascading effects on associated ecological processes in the periphyton community. These findings significantly enhance the understanding of the ecological impacts of nitrofurazone on marine periphytic ciliate communities, emphasizing the imperative for vigilant monitoring and regulation of veterinary antibiotics to protect marine ecosystem health and biodiversity. Further research is required to explore the long-term effects of nitrofurazone exposure and evaluate potential strategies to reduce the ecological repercussions of antibiotics in aquatic environments, with a particular focus on nitrofurazone.

The photic-aphotic divide is a strong ecological and evolutionary force determining the distribution of ciliates (Alveolata, Ciliophora) in the ocean.

The bulk of knowledge on marine ciliates is from shallow and/or sunlit waters. We studied ciliate diversity and distribution across epi- and mesopelagic oceanic waters, using DNA metabarcoding and phylogeny-based metrics. We analyzed sequences of the 18S rRNA gene (V4 region) from 369 samples collected at 12 depths (0-1,000 m) at the Bermuda Atlantic Time-series Study site of the Sargasso Sea (North Atlantic) monthly for 3 years. The comprehensive depth and temporal resolutions analyzed led to three main findings. First, there was a gradual but significant decrease in alpha-diversity (based on Faith's phylogenetic diversity index) from surface to 1,000-m waters. Second, multivariate analyses of beta-diversity (based on UniFrac distances) indicate that ciliate assemblages change significantly from photic to aphotic waters, with a switch from Oligotrichea to Oligohymenophorea prevalence. Third, phylogenetic placement of sequence variants and clade-level correlations (EPA-ng and GAPPA algorithms) show Oligotrichea, Litostomatea, Prostomatea, and Phyllopharyngea as anti-correlated with depth, while Oligohymenophorea (especially Apostomatia) have a direct relationship with depth. Two enigmatic environmental clades include either prevalent variants widely distributed in aphotic layers (the Oligohymenophorea OLIGO5) or subclades differentially distributed in photic versus aphotic waters (the Discotrichidae NASSO1). These results settle contradictory relationships between ciliate alpha-diversity and depth reported before, suggest functional changes in ciliate assemblages from photic to aphotic waters (with prevalence of algivory and mixotrophy versus omnivory and parasitism, respectively), and indicate that contemporary taxon distributions in the vertical profile have been strongly influenced by evolutionary processes. Integration of DNA sequences with organismal data (microscopy, functional experiments) and development of databases that link these sources of information remain as major tasks to better understand ciliate diversity, ecological roles, and evolution in the ocean.

FISHing for ciliates: Catalyzed reporter deposition fluorescence in situ hybridization for the detection of planktonic freshwater ciliates.

Planktonic ciliate species form multiple trophic guilds and are central components of freshwater food webs. Progress in molecular analytical tools has opened new insight into ciliate assemblages. However, high and variable 18S rDNA copy numbers, typical for ciliates, make reliable quantification by amplicon sequencing extremely difficult. For an exact determination of abundances, the classical morphology-based quantitative protargol staining is still the method of choice. Morphotype analyses, however, are time consuming and need specific taxonomic expertise. Catalyzed reporter deposition fluorescence in situ hybridization (CARD-FISH) may represent a promising tool for the analysis of planktonic ciliates by combining molecular identification with microscopic quantification. We tested the applicability of CARD-FISH using nine cultured ciliate species. Eight species- and three genus-specific oligonucleotide probes were designed based on their 18S rRNA genes. The CARD-FISH protocol was adapted and the specificity of probes was established. We subsequently examined the precision of quantitation by CARD-FISH on single cultures and mock assemblages. Successful tests on lake water samples proved that planktonic ciliates could be identified and quantified in field samples by CARD-FISH. Double hybridizations allowed studying interspecific predator prey interactions between two ciliate species. In summary, we demonstrate that CARD-FISH with species-specific probes can facilitate studies on the population dynamics of closely related, small sized or cryptic species at high sampling frequencies.

Four new species of Hysterocinetida (Protozoa: Ciliophora) from the digestive tract of earthworms collected to the lower Nyong estuary (South Coast, Cameroon)

Hysterocinetida are a large assemblage of ciliates characterized by sucker and buccal apparatus. They have a wide distribution in the digestive tract of oligochaete annelids of the genus Alma that inhabit hydromorphic parts of non-anthropic hydrosystems. On the basis of morphological, morphometric and infraciliature characteristics revealed by ammoniacal silver carbonate and 4',6-Diamidino-2-Phenyl Indole (DAPI) staining techniques, four forms were identified as new members belonging to three genera: Preptychostomum, Proptychostomum and Thurstonia. Preptychostomum donendaensis n. sp. is ovoid with a slightly narrowed anterior part and is commensal to the digestive tract of Alma emini and A. nilotica. There are 20-23 kineties on the lower side and 42-45 on the upper side. Proptychostomum gigas n. sp. is pear-shaped, with a narrowed anterior part and a wide posterior part (125-138 µm × 80-85 µm). There are 40 to 44 kineties, equally distributed on both sides of the ciliate. Its sucker is circular and measures 31-34 µm in diameter. The peristome follows the contour of the cell and the infundibulum has a turn of spire. Thurstonia emini n. sp. is identified in the digestive tract of A. emini. The cell is elongate, with the anterior pole narrower than the truncate posterior pole (90-115 µm × 40-55 µm). The kineties are equally distributed on both sides of the ciliate. The sucker is in the shape of an inverted U. Thurtonia nilotica n. sp. is commensal of the digestive tract of A. nilotica. The cell is pear-shaped, with the anterior part narrowed and the posterior part swollen (110-114 µm × 85-92 µm). 46 to 50 kineties are unevenly distributed on both sides of the ciliate. The sucker is shaped like an inverted lunar crescent.

Molecular insight reveals broad-scale spatial patterns in floodplain ciliate assemblages, whereas morphology reflects local environmental controls

One of the major goals in microbial ecology is to understand whether the empirical biogeographic patterns of macroorganisms also apply to microorganisms. Here, we used morphological data from live organisms, along with molecular data, to investigate the importance of spatial factors and environmental variables in influencing ciliate composition from floodplain lakes. Our main goal was to use 2 different approaches (morphological and molecular) to compare ciliate diversity and distribution patterns as well as to compare how these methods differ in their ability to detect distribution patterns and the roles of spatial and environmental factors that shape ciliate assemblages in the 4 largest floodplains in Brazil. Planktonic water samples were gathered from 33 lakes associated with 4 different river floodplain systems in Brazil. We analyzed ciliates in vivo and sequenced surface water DNA using a metabarcoding approach with general eukaryotic primers. We showed that the diversity of operational taxonomic units was much higher than that of morphospecies. Regardless of the method of identification, we found a consistent spatial assembly pattern of ciliate assemblages across the 4 floodplain systems. We also found that environmental filters had a stronger association with the morphological than with the molecular site-by-site dissimilarities. Meanwhile, biogeographic factors and the distance among sites limited the distribution of molecular-based composition, resulting in strong differences among the floodplain lakes analyzed. This finding suggests that ecological research and biomonitoring activities should find an equilibrium between morphological and molecular approaches because each approach provides unique insights.

Spatial Distribution of Ciliate Assemblages in a Shallow Floodplain Lake with an Anaerobic Zone

The spatial distribution of ciliate assemblages was studied in a shallow floodplain lake with a sharp division of space by oxygen conditions. The surface zone occupied by the “carpet” of Lemna trisulca and L. minor was characterized by a large daily amplitude of oxygen content with periodic exceeding of 100% of saturation; the underlying water layer was characterized by microaerobic conditions throughout most of the year, with seasonal deviations towards oxygen-free conditions (in winter and mid-summer) or increased oxygen content (before freezing and after ice melt); stable oxygen-free conditions were maintained in the bottom layer of water and at the bottom of the lake. There were 111 species of ciliated protozoa recorded in the lake. The ciliated protozoa were clearly structured and formed three almost non-overlapping assemblages in terms of species composition, which retained their isolation during all seasons of the year. On the basis of the analysis performed using the R indicspecies package, species of ciliated protozoa were identified as indicators of conditions with different oxygen regimes, which are determined by the level of organic pollution and the distribution of photosynthetic organisms.

High Dynamics of Ciliate Community Revealed via Short-Term, High-Frequency Sampling in a Subtropical Estuarine Ecosystem.

Ciliates are pivotal components of the marine microbial food web, exerting profound impacts on oceanic biogeochemical cycling. However, the temporal dynamics of ciliate assemblages on a short time scale in the highly fluctuating estuarine ecosystem remain largely unexplored. We studied changes in the ciliate community during a short time frame in the high salinity waters (>26) of a subtropical estuary. Ciliate abundance, biomass, size and oral diameter structure, and community composition fluctuated considerably and irregularly over a few days or even a few hours. Spearman correlations and the generalized linear model revealed that heterotrophic prokaryotes (HPs) and viral abundances drove the dynamics of ciliate abundance and biomass. The structural equation model further identified a major path from the high-fluorescence content virus (HFV) to HPs and then ciliates. Given the substantial correlation between salinity and HPs/HFV, we proposed that the response of HPs and HFV to salinity drives the dynamics of ciliate biomass. Additionally, the Mantel test showed that phytoplankton pigments such as Lutein and Neoxanthin, phosphate, and pigmented picoeukaryotes were key covariates of the ciliate community composition. This study demonstrated the highly changing patterns of ciliate assemblages and identified potential processes regulating ciliate biomass and community composition on short timescales in a subtropical, hydrographically complex estuary.

Deciphering phylogenetic relationships in class Karyorelictea (Protista, Ciliophora) based on updated multi-gene information with establishment of a new order Wilbertomorphida n. ord.

The class Karyorelictea, a unique assemblage of ciliates, is a key group in deciphering ciliate evolution history. However, the systematic relationships among members of this class remain poorly understood. Here we newly obtained eight small subunit (SSU) rDNA, 24 large subunit (LSU) rDNA, and 25 ITS1-5.8S-ITS2 sequences (covering 25 species, 10 genera and 4 out of 6 families) to analyze the phylogenetic relationships within Karyorelictea. Our results indicate that: (1) considering its unique morphology and early branching position in the SSU rDNA-based tree, the family Wilbertomorphidae represents a new taxon at order level, hence the new order Wilbertomorphida n. ord. is established; (2) all five families with available molecular information are monophyletic, as expected, and the orders Loxodida and Protostomatida show a closer relationship than with Protoheterotrichida; (3) in Trachelocercidae, the compound circumoral kineties is believed to be a plesiomorphic feature while the single circumoral kinety is synapomorphic; and (4) the freshwater genus Loxodes could be derived from the marine Remanella and both share most morphological features. Taken together, these muti-gene analyses provide further insights into the phylogeny of the diverse clades in Karyorelictea.

Ciliates in different types of pools in temperate, tropical, and polar climate zones – implications for climate change

Small water bodies are typically characterized by high diversity of various groups of microorganisms. Moreover, these ecosystems react very quickly to even the slightest climate changes (e.g. a temperature increase or water level fluctuations). Thus far, studies of planktonic ciliates in small water bodies having different origins and located in various climate zones have been scarce. Our study aimed to verify the following hypotheses: planktonic ciliate assemblages exhibit higher diversity in pools with higher concentrations of biogenic compounds; pools in warmer climates have higher biodiversity of planktonic ciliates than those in the polar climate zone; individual functional groups of ciliates demonstrate considerable diversity, both between individual pool types and between climate zones. The study was conducted in 21 small pools in temperate, tropical, and polar climate zones. While the type of pool clearly influenced the makeup of microbial communities, the influence of climate was stronger. The factors with the greatest influence on the occurrence of these microorganisms were temperature, total organic carbon, and nutrients. Our results show that in warmer climates the abundance of bacterivorous ciliates is higher, while that of mixotrophs is lower. This has consequences for modelling of climate change and assessment of its influence on the carbon cycle in small water body ecosystems.

Phylogenetic Position of Three Well-known Ciliates from the Controversial Order Loxocephalida Jankowski, 1980 (Scuticociliatia, Oligohymenophorea) and Urozona buetschlii (Schewiakoff, 1889) with Improved Morphological Descriptions

Loxocephalids represent a nonmonophyletic assemblage of oligohymenophorean ciliates with morphological features common to both scuticociliates (mode of ontogenesis) and hymenostomes (morphology of the oral structures). In phylogenetic analyses that include both nuclear and mitochondrial molecular markers, relationships among loxocephalid ciliates are still largely unresolved. With the aim to clarify the phylogeny of the controversial order Loxocephalida Jankowski, 1980, we provide the first 18S rRNA gene sequences of three morphologically well-described ciliates currently included in the order, namely Cinetochilum margaritaceum, Zitheron hovassei, and Zitheron muscorum comb. nov. We also provide the first 18S rRNA gene sequence for the monotypic genus Urozona. Here we present improved diagnoses for these species based on updated morphologic data. Our molecular data circumscribe the family Cinetochilidae Perty, 1852 and support the exclusion of genus Cinetochilides from family Cinetochilidae; therefore, we establish a family Cinetochilididae fam. nov. We consider Urozona as incertae sedis in Oligohymenophorea at the current state of knowledge.

Impact of El Niño-Southern Oscillation on plankton ciliates from a saline lowland river in South America

We analyzed the structure and dynamics of plankton ciliates from the Salado River, Buenos Aires (Argentina), during El Niño-Southern Oscillation events, to evaluate their response to alterations of certain environmental variables during these climatic phenomena. The basin exhibits intense land use at the headwaters and fluctuating salinity along the main channel, canals and tributaries. During El Niño (1997-1998), heavy rainfall caused high discharge, and low mean conductivity values were recorded along with low mean abundances in the entire basin. During neutral (2004-2005) and La Niña (2007-2009) events, the discharge was reduced, and mean conductivity was double the values recorded during El Niño, with maxima of mean abundances and diversity at the headwaters and the lower basin. The most abundant ciliates were tintinnids, heterotrichs and peritrichs. Species replacement was detected along the river system, mainly during the neutral event. At the headwaters and canals, abundance peaks of certain species were attributed to the strong human impact (nutrients and sewage inputs) and increased salinity, allowing the establishment of euryhaline ciliates. In the middle basin, freshwater species predominated, although less abundantly due to higher water flow. The mouth sector showed abundance peaks of an estuarine tintinnid. Temperature, conductivity and suspended solids were the most important variables driving ciliate structure. High discharge as a result of heavy rainfall during El Niño negatively influenced ciliate assemblages. In contrast, low water conditions during the other events allowed the development of diverse assemblages, increasing species number and abundance of particular species.

Spatial distribution of planktonic ciliates in the western Pacific Ocean: along the transect from Shenzhen (China) to Pohnpei (Micronesia).

Planktonic ciliates have been recognized as major consumers of nano- and picoplankton in pelagic ecosystems, playing pivotal roles in the transfer of matter and energy in the microbial loop. However, due to the difficulties in identification, the species composition of ciliate assemblages, especially for the small, fragile, and naked species that usually dominate the ciliate communities in the oceanic waters, remains largely unknown. In the present study, 22 stations along the transect from Shenzhen (China) to Pohnpei (Micronesia) were sampled for the enumeration of picoplankton and nanoflagellates. In addition, pigment analysis of major phytoplankton groups along with the measurements of environmental variables including temperature, salinity, and nutrients were also carried out. Ciliates were identified at species level using quantitative protargol stain to reveal the species composition and their distribution patterns from off-shore to open ocean. Ciliate abundance was positively correlated with phosphate, silicate, and pico-sized pigmented eukaryotes (PPEs), whereas the biomass was closely related with PPEs, heterotrophic nanoflagellates, and chlorophytes. The combination of silicate and pigmented nanoflagellates was identified as the major factor driving the ciliate community composition. The close relationship between silicate and ciliate abundance and community structure needs further validation based on more data collected from oceanic waters. Our study showed the necessity of using techniques that can reveal the community composition at higher taxonomic resolutions in future studies on ciliates.

Insights into the effects of harmful algal bloom on ecological quality status using body-size spectrum of biofilm-dwelling ciliates in marine ecosystems

The effects of two harmful algae Alexandrium tamarense and Gymnodinium catenatum on ecological quality status were identified using biofilm-dwelling ciliate assemblage as test organism communities. The body-size spectra of the test ciliates were observed at a gradient of cell concentrations of both algal species: 100 (control), followed by102, 103, 104 and 105 cells ml−1. The test ciliates showed clear variations in body-size spectra along the concentration gradients of both algal species. In terms of probability density, the ciliates generally peaked at low levels of algal concentrations (100–104 cells ml−1) in small size forms, followed by the forms with large sizes at the concentration of 105 cells ml−1 of both algal species. Bootstrapped-average analysis demonstrated a significant change in body-size spectrum when algal concentrations were higher than 104 cells ml−1. It is suggested that the body-size spectrum of the ciliates may be used to indicate the effects of harmful algal bloom.

Patterns in diversity and composition of the microbenthos of subarctic intertidal beaches with different morphodynamics

Diversity and composition of benthic diatom algae and ciliates were studied at several beaches along the White and Barents seas: from highly exposed, reflective beaches with coarse-grained sands to sheltered, dissipative silty-sandy flats. For diatoms, the epipelic to epipsammic species abundance ratio was significantly correlated with the beach index and mean particle size, while neither α-diversity measures nor mean cell length were related to beach properties. In contrast, most of the characteristics of ciliate assemblages (diversity, total abundance and biomass, mean individual weight and percentage of karyorelictids) demonstrated a strong correlation to beach properties, remaining low at exposed beaches but increasing sharply in more sheltered conditions. β-diversity did not correlate with beach properties for either diatoms or ciliates. We suggest that wave action and sediment properties are the main drivers controlling the diversity and composition of the intertidal microbenthos. Diatoms and ciliates, however, demonstrated divergent response to these factors. Epipelic and epipsammic diatoms exhibited 2 different strategies to adapt to their environments and therefore were complementarily distributed along the environmental gradient and compensated for each other in diversity. Most ciliates demonstrated a similar mode of habitat selection but differed in their degree of tolerance. Euryporal (including mesoporal) species were relatively tolerant to wave action and therefore occurred under a wide range of beach conditions, though their abundance and diversity were highest in fine, relatively stable sediments on sheltered beaches, whereas the specific interstitial (i.e. genuine microporal) species were mostly restricted to only these habitats.

Oxygen Gradients and Structure of the Ciliate Assemblages in Floodplain Lake

This paper presents the results of studies on the structure of the ciliate population in a freshwater lake. The classification of the ciliated communities based on the analysis of the distribution of ciliate population density in the lake along the oxygen gradients, taking into account their oxygen preferences, was proposed. It was shown that the distribution of ciliated protozoa in the space of a reservoir is determined not by such spatial units as the water column, bottom, and periphytal, but by the oxygen gradients. Four types of habitats with different oxygen regimes were distinguished: With stably high oxygen concentration, stably low oxygen concentration, stably oxygen-free conditions, and conditions with a high amplitude of diurnal oxygen variations. The location of these habitats in the space of the lake and their seasonal changes were determined. On the basis of the quantitative development of ciliate populations, zones of optima and tolerance ranges of some ciliate species in the oxygen gradient were established. The oxygen preferences were established for the species from four distinguished assemblages: Microoxyphilic, oxyphilic, euryoxyphilic, and anoxyphilic (anaerobic). The presence or the absence of a certain type of assemblage in the reservoirs depends solely on the parameters of the oxygen gradients. The diversity of the ciliated protozoa in water bodies also depends on the stability and diversity of the oxygen gradients.

Ciliate diversity and distribution across horizontal and vertical scales in the open ocean.

Ciliates are globally distributed eukaryotic organisms inhabiting virtually all environments on Earth. Although ciliates range from 10µm to a few millimetres in cell size, they are repeatedly reported in the pico-sized fraction (<2-3µm) of molecular surveys. Here, we used existing data sets (BioMarKs and Tara Oceans) with different size fractions to demonstrate that the ciliate pico-sized signal, probably derived from cell breakage during filtration, is informative and reliable to study marine ciliate biodiversity and biogeography. We then used sequences from the pico-eukaryotic fraction of two circumnavigation expeditions, Malaspina-2010 and Tara Oceans, to give insights into the taxonomic composition and horizontal and vertical distribution of ciliates in the global ocean. The results suggested a high homogeneity of ciliate communities along the ocean surface from temperate to tropical waters, with ciliate assemblages dominated by a few abundant and widely distributed taxa. Very few taxa were found in a single oceanic region, therefore suggesting a high level of ciliate cosmopolitanism in the global ocean. In vertical profiles, ciliates were detected up to 4,000m depth, and a clear vertical community structuring was observed. Our results provided evidence supporting ciliates as deeply integrated organisms in the deep-sea trophic web, where they may play a relevant role as symbionts of metazoans and grazers of prokaryotes and small eukaryotes in the water column and in aggregates.

Effectiveness of Photoprotective Strategies in Three Mixotrophic Planktonic Ciliate Species

Mixotrophic ciliate assemblages often prevail in summer in the surface layers of lakes. During this time, they are potentially exposed to damaging levels of incident solar ultraviolet radiation (UVR) and need efficient photoprotective mechanisms to minimize the damage. Herein, we tested the algal-bearing species of Pelagodileptus trachelioides, Stokesia vernalis, and Vorticella chlorellata for how they handled stress under exposure to the artificial sunlight spectrum (i.e., UV treatment), just photosynthetically active radiation (PAR), or in the dark (i.e., control). In addition to measurements of their survival, changes in behavior, shape, and whether dark or photoenzymatic repair (PER) mechanisms are present, we measured the concentration of UV-absorbing compounds (i.e., mycosporine-like amino acids). In contrast to the response in the PAR and dark treatments, sublethal effects were observed in all species when exposed to UVR. A wavelength-specific test for P. trachelioides revealed that UV-B was especially lethal. These results suggest that the photoprotective mechanisms found in these ciliates are not sufficient to allow for their survival directly at the surface and that, accordingly, they need to shift their position further down in the water column.