Microbial Food Webs in Marine Sediments. I. Trophic Interactions and Grazing Rates in Two Tidal Flat Communities

Abstract The role of grazing by marine sediment flagellates, ciliates, and meiobenthic animals in controlling production of their bacterial and diatom prey was investigated. Several novel or modified techniques were used to enumerate prey (bacteria and diatoms), measure bacterial production, quantify proto- and micrometazoan predators, and evaluate rates of bacterivory and herbivory. The results indicated that, in a temperate, marine intertidal flat composed of fine sand, colorless nanoflagellates, ciliates, and nematodes were the most important bacterivores. Together, these organisms were responsible for removing up to 53% of bacterial production, by grazing. The observed rates of bacterivory were high enough to hypothesize that periods of grazing control of bacterial production might occur regularly in similar habitats. Colorless microflagellates, ciliates, and nematodes had high rates of diatom consumption. The combined small diatom consumption rate was equivalent to 132% of diatom standing stock per day. Trophic interactions between diatoms and micro- and meiobenthos might be a factor limiting growth of small (around 10 μm) diatoms. In coarse sands of an open beach, all micrograzers except pigmented nanoflagellates were rare, whereas bacterial and diatom assemblages were rather abundant and active. In this type of sediment, the micrograzers were able to consume only a marginal percentage of bacterial production (<1%) and diatom standing stock (3.8%), thus playing a minor role in controlling the dynamics of their prey. Received: 11 June 1996; Accepted: 13 August 1996     

Channeling of bacterioplanktonic production toward phagotrophic flagellates and ciliates under different seasonal conditions in a river

The objective of this study was to analyze the flux of biomass through the communities of bacteria and phagotrophic protists in the cold and warm conditions occurring seasonally in Butrón River. Bacterial and heterotrophic protistan (flagellate and ciliate) abundance was determined by epifluorescence direct counts; protistan grazing on planktonic bacteria was measured from fluorescently labeled bacteria uptake rates; and the estimate of bacterial secondary production was obtained from [³H]thymidine incorporation rates. The abundance of bacterial, flagellate, and ciliate communities was similar during cold and warm situations. However, we observed that estimates of dynamic parameters, i.e., secondary bacterial production and protistan grazing, in both situations were noticeably different. In the warm situation, grazing rates of flagellates and ciliates (bacteria per protist per hour) were, respectively, 7 times and 18 times higher than those determined in the cold situation, and the grazing rates of the protistan communities (bacteria per protists present in 1 ml of water per hour) increased up to 5 times in the case of flagellates and 42 times in the case of ciliates. Estimates of bacterial secondary production were also higher during the warm situation, showing a ninefold increase. The percentage of bacterial production preyed upon by flagellates or ciliates was not significantly different between the two conditions. These results showed that in the different conditions of a system, the flux of biomass between the trophic levels may be quite different although this process may not be reflected in the abundance of each community of bacteria, flagellates, and ciliates. Offprint requests to: J. Iriberri.     

Microbial Community Structure and Dynamics in the Largest Natural French Lake (Lake Bourget)

We investigated the dynamics and diversity of heterotrophic bacteria, autotrophic and heterotrophic flagellates, and ciliates from March to July 2002 in the surface waters (0–50 m) of Lake Bourget. The heterotrophic bacteria consisted mainly of “small” cocci, but filaments (>2 μm), commonly considered to be grazing-resistant forms under increased nanoflagellate grazing, were also detected. These elongated cells mainly belonged to the Cytophaga-Flavobacterium (CF) cluster, and were most abundant during spring and early summer, when mixotrophic or heterotrophic flagellates were the main bacterial predators. The CF group strongly dominated fluorescent in situ hybridization–detected cells from March to June, whereas clear changes were observed in early summer when Beta-proteobacteria and Alpha-proteobacteria increased concomitantly with maximal protist grazing pressures. The analysis of protist community structure revealed that the flagellates consisted mainly of cryptomonad forms. The dynamics of Cryptomonas sp. and Dinobryon sp. suggested the potential importance of mixotrophs as consumers of bacteria. This point was verified by an experimental approach based on fluorescent microbeads to assess the potential grazing impact of all protist taxa in the epilimnion. From the results, three distinct periods in the functioning of the epilimnetic microbial loop were identified. In early spring, mixotrophic and heterotrophic flagellates constituted the main bacterivores, and were regulated by the availability of their resources mainly during April (phase 1). Once the “clear water phase” was established, the predation pressure of metazooplankton represented a strong top-down force on all microbial compartments. During this period only mixotrophic flagellates occasionally exerted a significant bacterivory pressure (phase 2). Finally, the early summer was characterized by the highest protozoan grazing impact and by a rapid shift in the carbon pathway transfer, with a fast change-over of the main predators contribution, i.e., mixotrophic, heterotrophic flagellates and ciliates in bacterial mortality. The high abundance of ciliates during this period was consistent with the high densities of resources (heterotrophic nanoflagellates, algae, bacteria) in deep layers containing the most chlorophyll. Bacteria, as ciliates, responded clearly to increasing phytoplankton abundance, and although bacterial grazing impact could vary largely, bacterial abundance seemed to be primarily bottom-up regulated (phase 3).     

Protistan bacterivory in an oligomesotrophic lake: Importance of attached ciliates and flagellates

Seasonal and depth variations of the abundance, biomass, and bacterivory of protozoa (heterotrophic and mixotrophic flagellates and ciliates) were determined during thermal stratification in an oligomesotrophic lake (Lake Pavin, France). Maximal densities of heterotrophic flagellates (1.9 × 10³ cells ml^–1) and ciliates (6.1 cells ml^–1) were found in the metalimnion. Pigmented flagellates dominated the flagellate biomass in the euphotic zone. Community composition of ciliated protists varied greatly with depth, and both the abundance and biomass of ciliates was dominated by oligotrichs. Heterotrophic flagellates dominated grazing, accounting for 84% of total protistan bacterivory. Maximal grazing impact of heterotrophic flagellates was 18.9 × 10⁶ bacteria 1^–1h^–1. On average, 62% of nonpigmented flagellates were found to ingest particles. Ciliates and mixotrophic flagellates averaged 13% and 3% of protistan bacterivory, respectively. Attached protozoa (ciliates and flagellates) were found to colonize the diatom Asterionella formosa. Attached bacterivores had higher ingestion rates than free bacterivorous protozoa and may account for 66% of total protozoa bacterivory. Our results indicated that even in low numbers, epibiotic protozoa may have a major grazing impact on free bacteria. Correspondence: C. Amblard.     

Size-selective grazing of coastal bacterioplankton by natural assemblages of pigmented flagellates,colorless flagellates,and ciliates

Fluorescently-labelled bacteria (FLB) were used to study the feeding strategies of a natural assemblage of estuarine protozoans and to examine whether the protozoan grazing could account for the in situ size structure of the bacterioplankton. The FLB, DTAF-stained enterococci, ranging in volume from 0.01 to 0.30 × 10^–1 µm³, were added to a natural planktonic assemblage at a density of 5.5% of the natural bacterioplankton. Initial densities (individuals ml^–1) were as follows: total natural bacteria, 2.2 × 10⁶; FLB, 1.2 × 10⁵; pigmented flagellates, 300; colorless flagellates, 250; and ciliates, 30. FLB consumption rates were determined by examining the contents of protozoan food vacuoles, and the long-term effect of grazing (over a period of 100 hours) was determined by monitoring the decline in the FLB density in experimental vessels. The average consumption rates of FLB by pigmented flagellates were similar to those by flagellates that lacked chloroplasts (0.9 and 0.6 FLB protozoan^–1 hour^–1, respectively). The ciliates consumed bacteria at an average rate that was 17-fold higher (per cell) than flagellates, and they displayed a greater preference for larger bacteria than did the flagellates. FLB of the mid-size classes (0.025–0.100 µm³) were heavily grazed by the entire protozoan assemblage; the smallest (<0.025 µm³) and the largest (>0.100 µm³) FLB escaped protozoan grazing. This had a profound effect on the resulting size distribution of FLB. At the end of a 100-hour incubation, the percentage of mid-size FLB (0.025 to 0.100 µm³) decreased 2.0–2.2-fold, while the percentage of the smallest and the largest FLB increased 2.0–2.5-fold. Resultant densities of FLB were consistent with initial clearance rates determined for the protozoan groups. The grazing rates of protozoans on FLB were species-specific; whereas some species consumed FLB, others did not demonstrate bacterivory. The results suggest that protozoan grazing has a major effect on the size distribution of coastal bacterioplankton. By selectively feeding on a particular size-class of bacteria, planktonic ciliates may consume 15–90% day^–1 of the standing stock of largest size classes of bacterioplankton. Thus, ciliates, which were present in low abundance in the field, could not balance the production of the entire bacterial community, but they may strongly influence the portion of the bacterial community represented by the largest bacterial class. The direct effect of flagellates (e.g., grazing) was limited to smaller bacteria.Offprint requests to: M. P. Shiaris.     

Viruses as Partners in Spring Bloom Microbial Trophodynamics

Population sizes of algae, bacteria, heterotrophic flagellates, and viruses were observed through the 1989 spring diatom bloom in Raunefjorden in western Norway. The culmination of the diatom bloom was followed by a peak in the concentration of bacteria and an increase in the concentration of heterotrophic flagellates, a pattern consistent with the concept of a food chain from photosynthetically produced organic material, through bacteria, to bacterivorous flagellates. The concentration of viruses varied through the spring bloom from 5 × 10⁵ in the prebloom situation to a maximum of 1.3 × 10⁷ viruses ml⁻¹ 1 week after the peak of the diatom bloom. Coinciding with the collapse in the diatom bloom, a succession of bacteria and viruses was observed in the mucous layer surrounding dead or senescent diatoms, with an estimated maximum of 23% of the total virus population attached to the diatoms. The dynamic behavior observed for the virus population rules out the possibility that it is dominated by inactive species, and the viruses are suggested to be active members of the microbial food web as agents causing lysis in parts of the bacterial population, diverting part of the bacterial production from the predatory food chain.     

On the structure and development of Arctic pack ice communities in Fram Strait: a multivariate approach

Summary The distribution of ice organisms was investigated in Fram Strait in May 1988 during the ARK V/1 expedition on RV Polarstern. Over a 3 week period the abundances of bacteria, diatoms, auto- and heterotrophic flagellates as well as various groups of meiofauna organisms were observed in the lowermost 30 cm of an ice floe. Data were obtained from three experimental fields under three different light regimes as a result of manipulations of the snow cover. The application of multivariate factor analysis on this time series data set resulted in the characterization of four succession stages of an Arctic sea ice community: 1) the diatom bottom assemblage, 2) the mixed autotrophic assemblage, 3) the mixed auto- and heterotrophic supra-bottom assemblage, and 4) the heterotrophic supra-bottom assemblage. The two most abundant meiofauna groups (Turbellaria, Ciliata) showed different preferences according to algal distribution. While turbellarians were most abundant in samples with mixed populations of diatoms and flagellates, ciliates reached their abundance maxima in samples dominated by diatoms, suggesting different prey selections. We have developed a model for the explanation of the spatial separation of auto- and heterotrophic organisms, highlighting the possible role of DOC production by ice algae and DOC transport with brine flow.     

Succession of Protists on Estuarine Aggregates

Abstract Colonization by and succession of bacteria and bacterivorous protists on laboratory-made aggregates were determined over a period of 14 days during winter and spring in 1997. Aggregates were generated from natural water from the limnetic zone of the Elbe Estuary using a tilting tube roller system. Within 1 h after the beginning of the experiments, macroaggregates started to form. Aggregates reached a maximum size of 1 mm with a tendency toward large sizes at the end of the experiment after the 10th day. On the first day, high bacterial densities of more than 10⁹ cells ml⁻¹ were detected within the aggregates. The abundances of flagellates and ciliates within aggregates were also two or three orders of magnitude higher than in the surrounding water. Densities of aggregate associated organisms are comparable to those occuring in sediments. The first protistan colonizers on the aggregates were small heterotrophic flagellates, such as choanoflagellates and small euglenids. Later, beginning on the 4th day, small sarcodines and ciliates became abundant. The most abundant ciliates associated with aggregates were small species of the Hypotrichia, Cyrtophorida, and Hymenostomata. After 9 days, large omnivorous and carnivorous ciliates, such as large members of the Hypotrichia and the Pleurostomatida, occurred. In spring, large heterotrophic flagellates and amebae also appeared at this time. These findings indicated the existence of a succession of protists on newly formed aggregates and a microbial food net within the aggregates based on bacterial production. Additionally, most of the species observed during this study were adapted for living on surfaces. In natural environments they are more common in benthic than in pelagic environments. For them, aggregates are havens in the water column comparable to sediment communities. Received: 7 January 2000; Accepted: 15 May 2000; Online Publication: 28 August 2000     

Dominance of microflagellates over diatoms in the Antarctic areas of deep vertical mixing and krill concentrations

Phytoplankton data obtained during six summer Polish expeditionsto the Antarctic Peninsula area, are compared with concurrentlyrecorded data on water column stabilities and krill abundance.The results show that flagellates (1.5–20 µm) arenumerically dominant over diatoms in the areas of deep verticalmixing and/or extensive krill concentrations. Of 102 stationsdominated by flagellates, 85 (83.3%) are located in a well mixedwater column (>100 m) and correspond to a mean krill densityof 15–346 t Nm^–2. In the same areas, estimated flagellatecarbon biomass exceeds diatom carbon. On the other hand, ofthe 40 stations dominated by diatoms, 36 (90%) are located inareas of increased water column stability (upper mixed layerof 10–50 m) and correspond to a low mean krill biomassof 0.34–4.6 t Nm^–2. Positive correlations of flagellateto diatom (F:D) cell number ratios with the depth of the uppermixed layer suggest light limitation of diatom growth and anincreased sinking rate of diatoms relative to flagellates inthe areas of deep vertical mixing. The relationship of the F:Dratio with krill abundance suggests that krill prefer feedingon diatoms and are less efficient in grazing particles of thesize of microflagellates (<20 µm). Flagellates exceeddiatoms in an unstable water column when the phytoplankton populationsare low; both algal groups increase in numbers with growingstability. The results provide field evidence that deep verticalmixing and krill grazing create conditions for the dominanceof flagellates over diatoms. Both factors acting together arelikely to suppress diatom blooms in the Antarctic.     

Phytoplankton species and abundance in response to eutrophication in coastal marine mesocosms

In a mesocosm nutrient enrichment experiment the species (orcategories) and abundances of diatoms, dinoflagellates, flagellates,monads and ciliates were identified and counted over a 16-monthperiod. Diatoms and ciliates increased with increasing nutrienttreatment while monads and flagellates, <10 µm in size,did not. By contrast, in the field diatoms sometimes appearedto decrease while small phytoplankton µ10 µm appearedto increase under eutrophic conditions. In the experiment, insome instances, grazing controlled abundances to low levelsin nutrient-enriched treatments. Self-shading by phytoplanktonlimited upper levels of abundance when nutrients were excessive.While nuisance species were occasionally present in variousnutrient treatments, the intensity and frequency of their presencedid not tend to increase with nutrient treatment. Generallyspecies (or categories) did not appear to change with nutrienttreatment.     

Seasonal and successional influences on bacterial community composition exceed that of protozoan grazing in river biofilms

The effects of protozoa (heterotrophic flagellates and ciliates) on the morphology and community composition of bacterial biofilms were tested under natural background conditions by applying size fractionation in a river bypass system. Confocal laser scanning microscopy (CLSM) was used to monitor the morphological structure of the biofilm, and fingerprinting methods (single-stranded conformation polymorphism [SSCP] and denaturing gradient gel electrophoresis [DGGE]) were utilized to assess changes in bacterial community composition. Season and internal population dynamics had a greater influence on the bacterial biofilm than the presence of protozoa. Within this general framework, bacterial area coverage and microcolony abundance were nevertheless enhanced by the presence of ciliates (but not by the presence of flagellates). We also found that the richness of bacterial operational taxonomic units was much higher in planktonic founder communities than in the ones establishing the biofilm. Within the first 2 h of colonization of an empty substrate by bacteria, the presence of flagellates additionally altered their biofilm community composition. As the biofilms matured, the number of bacterial operational taxonomic units increased when flagellates were present in high abundances. The additional presence of ciliates tended to at first reduce (days 2 to 7) and later increase (days 14 to 29) bacterial operational taxonomic unit richness. Altogether, the response of the bacterial community to protozoan grazing pressure was small compared to that reported in planktonic studies, but our findings contradict the assumption of a general grazing resistance of bacterial biofilms toward protozoa.     

Seasonal plankton dynamics in a Mediterranean hypersaline coastal lagoon: the Mar Menor

The seasonal distribution of plankton in a Mediterranean hypersalinecoastal lagoon has been studied through a dataset, comprisingthe taxonomic composition and the size–abundance distributionof both phyto- and zooplankton, measured by image analysis techniquesduring a one-year time series of weekly samplings. The studiedorganisms ranged from small nanoplanktonic heterotrophic flagellates(2 µm diameter) to fish larvae (>2 µm). The phytoplanktonannual succession was characterized by a winter period dominatedby Rhodomonas spp. and Cryptomonas spp. with Cyclotella spp.as the main diatom represented, a spring phase where diatoms(mainly Cyclotella) were the dominant group with some monospecificblooms of other diatoms (mainly of Chaetoceros sp.), a summerphase characterized by diatoms with blooms of Niztschia closterium,and a post-summer phase where dinoflagellates increased withpeaks of Ceratium furca. High densities of the microbial foodweb elements, flagellates and ciliates, indicate the importanceof the microbial loop in the ecosystem. Meroplankton contributedwidely to the seasonal character of the zooplankton distribution.Copepods, represented by Oithona nana, Centropages ponticusand Acartia spp. (mainly latisetosa), remained relatively constantthroughout the year, exhibiting a lower density in the warmerwater period (July–September). At the end of the samplingperiod, a massive proliferation of copepods (>1000 ind l^–1), mainly due to O. nana, took place. The autotrophsto heterotrophs biovolume ratio (A:H) remained lower than 1throughout the year except when, occasionally, large phytoplanktoncells bloomed. Persistent very low values of A:H suggest thatadditional sources of energy, such as the microbial loop ordetrital pathways, would be needed to sustain the high heterotrophicbiovolume found in the lagoon.     

Abundance and Biomass of Heterotrophic Flagellates, and Factors Controlling Their Abundance and Distribution in Sediments of Botany Bay

The abundance and biomass of heterotrophic flagellates were estimated monthly in sediments of Botany Bay during March 1999-February 2000. The annual abundance and biomass were in the ranges of 0.46-4.70 x 10(5) cells/cm(3) and of 0.30-8.61 micro g C/cm(3), respectively. The majority of heterotrophic flagellates (93-100%) were less than 10 mm in length and few flagellates were larger than 10 mm. Of the total microbial carbon biomass, heterotrophic flagellates made up about 5% (but at times up to 35%). The contribution of heterotrophic flagellates varied from month to month, and among the sites. The abundance of heterotrophic flagellates was negatively correlated with sediment grain size and positively correlated with the abundance of bacteria, algae (autotrophic flagellates and diatoms), and their probable grazers. A best subsets regression analysis showed that bacterial and algal abundance are the most important factors controlling the abundance of heterotrophic flagellates. When the previously reported grazing rates on bacteria were applied, heterotrophic flagellates would consume a maximum of 64% of bacterial standing stock daily in Botany Bay, suggesting that heterotrophic flagellates are important as bacterivores. However, the importance of heterotrophic flagellate grazing probably varies significantly among the sites and from month to month.     

Microbial Response of a Freshwater Benthic Community to a Simulated Diatom Sedimentation Event: Interactive Effects of Benthic Fauna

Abstract The response of a sediment microbial assemblage to a pulse of diatoms was studied over 36 days by measuring bacterial activity and biomass, ATP concentration, and overall community respiration in laboratory microcosms. Also, the contribution of macrofaunal chironomids to the decomposition of settling diatoms in benthic communities, and the relative importance of benthic meiofauna in community metabolism, were determined. The addition of diatoms resulted in an immediate response by sediment bacteria, with higher bacterial production recorded after only 2 h, and a more than tenfold increase within one day. The rapid response by sediment bacteria was accompanied by relatively high initial concentrations of dissolved organic carbon. In treatments receiving diatoms, higher bacterial production was sustained throughout the experiment. Surprisingly, neither these elevated production estimates, nor the starvation of controls affected bacterial abundance. Mean bacterial cell volume, however, was markedly affected by the addition of diatoms. Combining community respiration measurements and bacterial production estimates showed that growth efficiencies for sediment bacteria ranged from 14.6 to 34.5%. The contribution of ambient meiozoobenthos to carbon metabolism was less than 1%. Carbon budgets showed that 1.3 mg C was cooxidized along with 4.3 mg added diatom C. Sediment reworking by Chironomus larvae initially enhanced bacterial production, but the presence of Chironomus resulted in lower bacterial production estimates after 16 and 36 days. This was interpreted as a result of faster decomposition of diatoms in treatments with chironomids, which was validated by a faster decline of ATP and chlorophyll a in the sediment. Our results indicate that Chironomus larvae compete with sediment bacteria for available organic substrates. Received: 11 June 1996; Accepted: 13 August 1996     

Protozoan Bacterivory in the Ice and the Water Column of a Cold Temperate Lagoon

Abstract Bacterial abundance and bacterivorous protist abundance and activity were examined in ice-brine and water column communities of a cold temperate Japanese lagoon (Saroma-Ko Lagoon, Hokkaido, 44°N, 144°E), during the late winter phase of ice community development (February–March 1992). Bacterial abundance averaged 6 and 1 × 10⁵ cells ml⁻¹ in the ice-brine and plankton samples, respectively, and generally decreased during the sampling period. Bacterivorous protists, identified based on direct observation of short-term (<1 h) ingested fluorescently labeled bacteria (FLB) in their food vacuoles, were largely dominated by flagellates, mainly cryothecomonad-type and chrysomonad-like cells and small dinoflagellates of the genus Gymnodinium. Bacterivorous ciliates included mainly the prostomatid Urotricha sp., the scuticociliates Uronema and Cyclidium, the choreotrichs Lohmaniella oviformis and Strobilidium, and the hypotrich Euplotes sp. Protist abundance averaged 4 × 10³ and 8.1 cells ml⁻¹ in the ice-brine and 0.3 × 10³ and 1.2 cells ml⁻¹ in the plankton, for flagellates and ciliates, respectively. In contrast to bacteria, the abundance of protists generally increased throughout the sampling period, indicating predator–prey interactions. Protistan bacterivory, measured from the rate of FLB disappearance over 24 h, averaged 36% (ice) and 24% (plankton) of bacterial standing stock and exhibited the same seasonal pattern as for protist abundance. The calculated specific clearance (range, 2–67 nl protozoa⁻¹ h⁻¹) and ingestion (<1–26 particles protozoa⁻¹ h⁻¹) rates were likely to be minimal estimates and grazing impact may have been higher on occasion. Indications for the dependence of ``bacterivorous protists' on nonbacterial food items were also provided. Although alternative sources of bacterial loss are likely to be of importance, this study provides evidence for the potential of protozoan assemblages as bacterial grazers in both sea ice-brine biota and water column at the southern limit of sea ice in the northern hemisphere. Received: 30 July 1998; Accepted: 18 November 1998     

Bacterivory and herbivory: Key roles of phagotrophic protists in pelagic food webs

Research on microbial loop organisms, heterotrophic bacteria and phagotrophic protists, has been stimulated in large measure by Pomeroy's seminal paper published in BioScience in 1974. We now know that a significant fate of bacterioplankton production is grazing by < 20-µm-sized flagellates. By selectively grazing larger, more rapidly growing and dividing cells in the bacterioplankton assemblage, bacterivores may be directly cropping bacterial production rather than simply the standing stock of bacterial cells. Protistan herbivory, however, is likely to be a more significant pathway of carbon flow in pelagic food webs than is bacterivory. Herbivores include both < 20-µm flagellates as well as > 20-µm ciliates and heterotrophic dinoflagellates in the microzooplankton. Protists can grow as fast as, or faster than their phytoplankton prey. Phototrophic cells grazed by protists range from bacterial-sized prochlorophytes to large diatom chains (which are preyed upon by extracellularly-feeding dinoflagellates). Recent estimates of microzooplankton herbivory in various parts of the sea suggest that protists routinely consume from 25 to 100% of daily phytoplankton production, even in diatom-dominated upwelling blooms. Phagotrophic protists should be viewed as a dominant biotic control of both bacteria and of phytoplankton in the sea.     

Direct and indirect influences of crustacean zooplankton on bacterioplankton of Lake Constance

Herbivorous crustacean zooplankton may influence bacterial populations of lakes directly by grazing on them or indirectly by grazing on algae. In Lake Constance a regularly observed decrease of bacterial density during periods of high abundance of cladocerans (clearwater phase) indicated bacterial grazing losses. However, cladoceran grazing on bacteria appeared to be less efficient than on algae. Moreover, cladocera reduced grazing pressure on bacteria by grazing on bacterivorous flagellates. Additionally, a shift of bacterial composition from an originally higher percentage of filamentous and aggregate growth forms towards a population of homogenously distributed small single celled bacteria was observed regularly at the beginning of the clearwater phase. Transient increases of bacterial abundance and productivity coinciding with the increase of cladocera at the end of the algal spring bloom were interpreted as field indications of indirect bacteria-zooplankton interactions due to crustacean grazing on phytoplankton. The release of organic carbon during grazing of crustacea on algae was considered as explanation for the observed stimulation of bacterial populations. Thereby, additional, otherwise inaccessible algal carbon would be made available to bacteria by zooplankton. Experimental support for this hypothesis was given by showing that bacteria were able to respond to crustacean grazing on algae by enhanced growth and activities. The possible impact of these direct and indirect crustacea-bacteria interactions on the abundance, activity and composition of bacterioplankton as well as on the structure and function of the total planktonic community is discussed.     

The importance of bacterivorous protists in the decomposition of stream leaf litter

1. Allochthonous organic matter, in the form of senesced leaves, is a major source of carbon supporting detrital food webs. While studies have documented the role of bacteria and fungi in the decomposition of leaf litter, little information is available regarding the role of protists in the decomposition process. 2. We tested the hypothesis that the presence of stream‐dwelling bacterivorous protists leads to an increased rate of leaf decomposition through grazing pressure on bacteria. We isolated live protists from decomposing leaves collected in a stream in Northern Virginia, U.S.A. (Goose Creek) and established laboratory cultures of common bacterivorous protists. 3. Recently senesced leaves from the field were used in laboratory microcosm experiments to determine if the rate of litter decomposition differed between four treatments: bacteria only, bacteria + flagellates, bacteria + flagellates + ciliates, autoclaved stream water (control). We determined the dry weight of leaf remaining, bacterial abundance, flagellate abundance and ciliate abundance for each replicate on days 0, 7, 14, 30, 60 and 120. 4. The rate of leaf decomposition was significantly higher in treatments with protists than without and bacterial abundance declined in protist treatments compared with bacteria only treatment. Weight loss in the presence of flagellates was three to four times higher when protists were present compared with treatments with bacteria alone. These results provide experimental evidence that protists could play a significant role in the detrital processes of streams.     

Possible food chain relationships between bacterioplankton,protozoans, and cladocerans in a reservoir

Ingestion of fluorescent particles by natural protozoan assemblage was studied in the Řimov Reservoir (Southern Bohemia) from April to October, 1987. Attached and free-living bacterial abundance, proportion of active bacteria, density of suspended particles and biomass of cladocerans were also monitored. Heterotrophic nanoflagellates (HNF; 5–12.8 10²ml⁻¹) were the dominant bacterial micrograzers during the spring period and consumed 3 to 9% of the total bacteria per day. After the spring phytoplankton bloom maximum densities of suspended particles and attached bacteria (up to 28% of the total counts) were found. Development of cladocerans in May sharply decreased the proportion of attached bacteria and kept them below 5% of the total counts. All the studied components of plankton except Cladocera decreased during the clearwater phase. The most significant drop was observed in the numbers of protozoans, and they were negligible for bacterial elimination. Bacterial losses during that time apparently were due to cladoceran grazing. During the summer period, ciliates (15–142 ml⁻¹) were mostly dominant micrograzers, and protozoan community grazing increased up to 21% of bacterial standing stock per day. The proportion of active bacteria was strongly correlated with protozoan grazing (r=0.83).     

Ciliate grazing on <Emphasis Type="Italic">Nitrosomonas europaea</Emphasis> and <Emphasis Type="Italic">Nitrospira</Emphasis><Emphasis Type="Italic">moscoviensis</Emphasis>: Is selectivity a factor for the nitrogen cycle in natural aquatic systems?

Ciliated protists are important predators of bacteria in many aquatic habitats, including sediments. Since, many biochemical transformations within the nitrogen cycle are performed by bacteria, ciliates could have an indirect impact on the nitrogen cycle through selective grazing on nitrogen-transforming bacteria. As a case study, we examined ciliate grazing on nitrifying bacteria of the genera Nitrosomonas and Nitrospira. All experiments were designed as in vitro-experiments with cultures of different bacteria and ciliate species. The nitrifying bacteria used in our experiments were Nitrosomonas europaea [Winogradsky 1892] and Nitrospira moscoviensis [Ehrich 2001]. The ciliates comprised of four species that are known as efficient bacterivores and common members of the protist community in aquatic systems: Paramecium aurelia [Müller 1773], Euplotes octocarinatus [Carter 1972], Tetrahymena pyriformis [Ehrenberg 1830] and Cyclidium glaucoma [Müller 1786]. Our experimental approach, using a combination of DAPI and FISH staining, was successful in allowing the observation of ingestion of specific bacteria and their detection within ciliate food vacuoles. However, the ciliates in this study showed no significant selective grazing. No food preferences for a any bacterial taxon or any size class or morphotype were detected. Correlation with time between ciliate abundance and bacterial abundance or biovolume, using log transformed growth rates of ciliates and bacteria, showed no significant results. On the bacterial side, neither an active defence mechanism of the nitrifying bacteria against ciliate grazing, such as changes in morphology, nor competition for resources were observed. These results suggest that in our in vitro-experiments grazing by ciliates has no influence on abundance and growth of nitrifying bacteria and nitrification.