Microbial Food Webs in Marine Sediments. II. Seasonal Changes in Trophic Interactions in a Sandy Tidal Flat Community

Abstract The role of grazing by marine sediment flagellates, ciliates, and meiobenthic animals in controlling production of their bacterial and diatom prey was investigated. At six selected time points, over the year, bacterial production and diatom standing stock were compared to grazing pressure exercised by proto- and micrometazoan consumers. The intensity of prey-predator relations showed pronounced yearly dynamics in which two stages could be distinguished. During the first phase, from the end of winter to mid-summer, the consumption of diatoms gradually increased, with possible overgrazing at the end of the period. This was followed by a collapse of diatom abundance, to the winter level. During the first stage, no appreciable bacterial consumption was observed in spite of the high abundance and production of bacteria. The second stage started in mid-summer and continued through the fall. During this period, the grazing on bacteria increased and reached the year's maximum. For at least a brief period (October), micrograzers removed the majority of bacterial production. In contrast, herbivory stayed at the year's lowest level, and diatoms appeared to be controlled by factors other than grazing. The observed ingestion rates seem to support the apparent energy requirements of flagellates and some ciliates (scuticociliates and hypotrichids). Other ciliates (pleurostomatids and karyorelictids) could not subsist on the observed diet and might have to complement it with other energy sources, possibly via dissolved organic matter absorption. Received: 27 August 1996; Accepted: 22 January 1997     

Rates of Microbenthic and Meiobenthic Bacterivory in a Temperate Muddy Tidal Flat Community

Rates of bacterivory in micro- and meiobenthic species were determined by an improved technique in a muddy tidal flat community in Boston Harbor, Mass. The predominant grazers of bacteria were identified, and their rates of grazing were measured in the top 1 cm of the sediment. Grazing rates were measured by a fluorescence-labeled bacteria (FLB) technique. A mixture of two Enterococcus spp. isolates and two isolates of Escherichia coli were prepared as FLB, and they were added to intact sediment cores by replacing the pore water in the upper centimeter of the core. A standard FLB procedure was modified by filtering sediment dilutions onto cellulose membrane filters and processing the filters to render them optically transparent while preserving the physical integrity of the micro- and meiobenthic organisms. Thus, it was possible, on the same microscopic field, to switch from light microscopy for identification of grazers to epifluorescence microscopy for counting FLB present in the gut contents of the same grazers. The majority of benthic organisms present in these sediments consumed FLB, but their consumption rates varied widely. Two ciliate species, a Prorodon sp. and a Chlamidodon sp., and a nematode, a Metoncholaimus sp., consumed fluorescence-labeled coliforms at the highest rates, 126 to 169 FLB per individual per h. Other ciliates and nematodes, as well as microflagellates and harpacticoid copepods, consumed fluorescence-labeled coliforms at lower rates, 1.2 to 26 FLB per individual per h. Foraminiferans and gastrotriches did not contain FLB. Some ciliate grazers discriminated between enterococci and coliforms, consuming the rod-shaped fluorescence-labeled coliforms at 74- to 155-fold-higher rates than did the coccus-shaped fluorescence-labeled enterococci. Other ciliates did not select between fluorescence-labeled enterococci and fluorescence-labeled coliforms. The high rates of bacterivory by some ciliates and nematodes indicated intensive grazing. However, at their low extant densities, the grazers consumed only a small portion of the bacterial standing stock. Major bacterial grazers, e.g., microflagellates, ciliates, and nematodes, could potentially consume, per day, only 0.2, 0.1, and 0.03%, respectively, of the bacterial standing stock (7.5 × 10⁸ bacteria per cm³).     

Benthic Bacterial Production and Protozoan Predation in a Silty Freshwater Environment

The interrelation of heterotrophic bacteria with bacterivorous protists has been widely studied in pelagic environments, but data on benthic habitats, especially in freshwater systems, are still scarce. We present a seasonal study focusing on bacterivory by heterotrophic nanoflagellates (HNF) and ciliates in the silty sediment of a temperate macrophyte-dominated oxbow lake. From January 2001 to February 2002 we monitored the standing stock of bacteria and protozoa, bacterial secondary production (BSP, ³H-thymidine, and ¹⁴C-leucine incorporation), and grazing rates of HNF and ciliates on bacteria (FLB uptake) in the oxic sediment of the investigated system. BSP ranged from 470 to 4050 µg C L^–1 wet sediment h^–1. The bacterial compartment turned out to be highly dynamic, indicated by population doubling times (0.6–10.0 d), which were comparable to those in the water column of the investigated system. Yet, the control mechanisms acting upon the bacterial population led to a relative constancy of bacterial standing stock during a year. Ingestion rates of protozoan grazers were 0–20.0 bacteria HNF^–1 h^–1 and 0–97.6 bacteria ciliate^–1 h^–1. HNF and ciliates together cropped 0–14 (mean 4)% of BSP, indicating that they did not significantly contribute to benthic bacterial mortality during any period of the year. The low impact of protozoan grazing was due to the low numbers of HNF and ciliates in relation to bacteria (1.8–3.5 × 10⁴ bacteria HNF^–1, 0.9–3.1 × 10⁶ bacteria ciliate^–1). Thus, grazing by HNF and ciliates could be ruled out as a parameter regulating bacterial standing stock or production in the sediment of the investigated system, but the factors responsible for the limitation of benthic protistan densities and the fate of benthic BSP remained unclear.     

Effects of Deposit-Feeding Macrofauna on Benthic Bacteria,Viruses, and Protozoa in a Silty Freshwater Sediment

In microcosm experiments, we simultaneously tested the effects of increased numbers of deposit-feeding macrofauna (chironomids, oligochaetes and cladocerans) on the standing stock, activities and interactions of heterotrophic bacteria, viruses, and bacterivorous protozoa (heterotrophic nanoflagellates and ciliates) in the aerobic layer of a silty littoral freshwater sediment. On average, bacterial secondary production was stimulated between 11 and 29% by all macrofaunal groups compared to control experiments without macrofauna addition. Bacterial standing stock increased significantly by 8 and 13% in case of chironomids and cladocerans, respectively. Oligochaetes and chironomids produced significant negative effects on viral abundance while the results with cladocerans were inconsistent. The addition of oligochaetes and chironomids resulted in a significant decrease by on average 68 and 32% of viral decay rates, respectively, used as a measure of viral production. The calculated contribution of virus-induced lysis to benthic bacterial mortality was low, with 2.8 to 11.8% of bacterial secondary production, and decreased by 39 to 81% after the addition of macrofauna compared to the control. The abundances of heterotrophic nanoflagellates were significantly reduced by 20% by all tested macrofauna groups, while ciliates showed inconsistent results. The importance of heterotrophic nanoflagellate grazing on benthic bacteria was very low (<1% of bacterial secondary production) and was further reduced by elevated numbers of macrofauna. Thus, the selected deposit feeding macrofauna groups seem to have several direct and indirect and partly antagonistic effects on the benthic bacterial compartment through the enhancement of bacterial production and the reduction of virus-induced cell lysis and protozoan grazing.     

Bacterioplankton production and protozoan bacterivory in a mesotrophic reservoir

Protozoan bacterivory [via uptake of fluorescently labelledbacteria (FLB)] and production of bacteria ([³H]thymidine assay)were simultaneously measured in the mesotrophic ímovReservoir (Southern Bohemia) from April to November, 1988. Heterotrophicnanoflagellates (HNF) were mostly responsible for a greaterfraction of protozoan bacterivory during the spring period.From 10 to 23% of bacterial production was grazed daily withthe only exception of the spring peak of ciliate abundance (upto 60%). Protozoans decreased significantly during the clearwaterphase (ciliates disappeared), and thus their role in bacterivorywas negligible. Through the summer-fall period ciliates, notHNF, were the most important bacterial micrograzers. Protozoancommunity grazing balanced or even exceeded the daily bacterialproduction in August and September. Alternate fates of bacterialproduction besides protozoan grazing during the spring periodare discussed.     

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).     

Ciliates are the Dominant Grazers on Pico- and Nanoplankton in a Shallow,Naturally Highly Eutrophic Lake

Abundance and biomass of the microbial loop members [bacteria, heterotrophic nanoflagellates (HNF), and ciliates] were seasonally measured in the naturally eutrophic and shallow (2.8 mean depth) Lake Võrtsjärv, which has a large open surface area (average 270 km²) and highly turbid water (Secchi depth <1 m). Grazing rates (filter feeding rates) on 0.5-, 3-, and 6-μm-diameter particles were measured to estimate pico- and nanoplankton grazing (filter feeding) by micro- and metazooplankton. Among grazers, HNF had a low abundance (<50 cells mL^?1) and, due to their low specific filtering rates, they only grazed a minor fraction of the bacterioplankton (≤4.2% of total grazing). Ciliates were relatively abundant (≤158 cells mL^?1) and, considering their high specific feeding rates, were able to graze more than 100% of the bacterial biomass production in the open part of the lake, whereas the average daily grazing accounted for 9.3% of the bacterial standing stock. Ciliates were potentially important grazers of nanoplanktonic organisms (on average, approximately 20% of the standing stock of 3-μm-size particles was grazed daily). Metazooplankton grazed a minor part of the bacterioplankton, accounting for only 0.1% of standing stock of bacteria. Grazing on nanoplankton (3–6 μm) by metazooplankton was higher (0.4% of standing stock). The hypothesis is proposed that ciliates dominate due to a lack of top–down regulation by predators, and HNF have a low abundance due to strong grazing pressure by ciliates.     

The composition of particulate organic matter and biomass in the Peruvian upwelling region during ICANE 1977 (Nov. 14 - Dec. 2)

This paper presents the results of plankton studies in the Peruvianupwelling region off Chimbote from November - December, 1977,during the Investigación Cooperativa de la Anchovetay su Ecosistema (ICANE). Primary productivity, respiratory ETSactivity, composition of particulate organic matter, and microplanktoncell numbers were determined. Phytoplankton growth, and bacterialand ciliate carbon-uptake rates were computed from cell counts. Inshore waters were dominated by diatoms and were more productivethan offshore waters which were dominated by dinoflagellatesand ciliates. Particulate primary production averaged 5.26 ±5.24 g C m^–2d^–1, and the POC standing stock was7.75 ± 2.74 g C m^–2 for the euphotic layer of 7shelf stations. On the shelf, microplankton respiration rateswere higher in plankton populations dominated by dinoflagellatesand ciliates (47% of particulate primary production per 24 h)than those in diatom dominated populations (11%, respectively).The diatom populations, which were dominated by Chaetocerosspecies, varied in their ecophysiological properties (assimilationnumbers, proportion of water soluble carbohydrate, and protein/nitrogenratios). The relationships between these variations and growthconditions were investigated. A 40 h time-series station revealedpatchiness which was superimposed on physiological changes ofthe plankters. Bacterial numbers of 2 x 10⁶ cells/ml were foundin the euphotic layer corresponding to approximately 17 µgC/l bacterial biomass (or 6% of the POC standing stock). Ciliatebiomass (Lohmanniella oviformis was the dominating species)ranging from 2 to 9% of the POC standing stock were found evenin diatom dominated populations. From a rough carbon balancefor the euphotic layer it was deduced that in diatom dominatedpopulations, 36–77% of particulate primary productionwas potentially available to adult anchoveta grazing.     

Nanoflagellate predation on auto- and heterotrophic picoplankton in the oligotrophic Mediterranean Sea

Dynamics of autotrophic and heterotrophic prokaryotes and theirconsumption by nanoflagellates were studied in the euphoticzone at nine stations located from the Levantine Basin (34°E)to the Balearic sea (5°E) in June 1999. Bacterial biomassconstituted the largest proportion of living biomass at allstations. Integrated bacterial production at the furthest eaststation, was sixfold lower than integrated bacterial productionat the furthest west (13 and 75 mg C m^-2 d^-1 respectively).Estimated heterotrophic nanoflagellate bacterivory accountedfor 45–87% of bacterial production. Small protists (<3µm) dominated the bacterivore assemblage and accountedfor more than 90% of the heterotrophic bacterial consumption.Our results indicated that there was no negative selection againstSynechococcus and that both picoplankton groups were grazedaccording to their standing stocks. An estimated consumptionof Synechococcus derived from food vacuole content analysisof nanoflagellates revealed that they consumed from 0.5 to 45%(mean 13%) of Synechococcus stock per day. These data are amongthe first documenting the relative grazing impact of heterotrophicnanoflagellates on bacteria and Synechococcus in situ. Assumingthat there was no selection for or against Prochlorococcus,heterotrophic nanoflagellates could ingest from 1.4 to 21% (mean6%) of Prochlorococcus stock per day. The amount of organiccarbon obtained by heterotrophic nanoflagellates from photosyntheticprokaryotes represented 27% of the total amount of carbon obtainedfrom total prokaryotes     

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.     

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     

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.     

Effect of Protistan Grazing on the Frequency of Dividing Cells in Bacterioplankton Assemblages

Grazing by phagotrophic flagellates and ciliates is a major source of mortality for bacterioplankton in both marine and freshwater systems. Recent studies have demonstrated a positive relationship between clearance rate and prey size for bacterivorous protists. We tested the idea that, by selectively grazing the larger (more actively growing or dividing) cells in a bacterial assemblage, protists control bacterial standing stock abundances by directly cropping bacterial production. Samples of estuarine water were passed through 0.8-μm-pore-size filters (bacteria only) or 20-μm-mesh screens (bacteria and bacterivorous protists) and placed in dialysis tubing suspended in 7 liters of unfiltered water. Changes in total bacterial biovolume per milliliter (bacterial biomass), frequency of dividing cells (FDC), and average per cell biovolume were followed over a period of 24 h. In three experiments, the FDC increased more rapidly and attained higher values in water passed through 0.8-μm-pore-size filters (average, 5.1 to 8.9%; maximum, 15.5%) compared with FDC values in water passed through 20-μm-mesh screens (average, 2.7 to 5.3%; maximum, 6.7%). Increases in bacterial biomass per milliliter lagged behind increases in FDC by about 4 to 6 h. Grazed bacterial assemblages were characterized by lower total biomasses and smaller average cell sizes compared with those of cells in nongrazed assemblages. We conclude that bacterivorous protists control bacterial standing stock abundances partly by preferentially removing dividing cells. Selective grazing of the more actively growing cells may also explain, in part, the ability of slow-growing cells to persist in bacterioplankton assemblages.     

Protist herbivory: a key pathway in the pelagic food web of Lake Tanganyika

Herbivory and bacterivory by phagotrophic protists were estimated in the southern basin of the oligotrophic Lake Tanganyika at different seasons (in the rainy season in February?CMarch 2007 and in the dry season in July?CAugust 2006 and September 2007), using two independent methods: the selective inhibitor technique for assessing community grazing on picocyanobacteria (PCya) and fluorescently labelled bacteria (FLB) and Synechococcus (FLA) to estimate bacterivory and herbivory by phagotrophic nanoflagellates (NF) and ciliates. Protistan grazing impact on both heterotrophic bacteria and PCya was mainly due to NF, which contributed up to 96% of the microbial grazing. There was a clear selection of FLA by protists. PCya represented the main carbon source for both flagellates and ciliates in the mixolimnion, accounting for an average of 83% of the total carbon obtained from the ingestion of picoplanktonic organisms. Protists were the main consumers of particulate primary production (46?C74% depending on season). Significant seasonal variation of grazing rates (0.011?C0.041?h^?1) was found, chiefly following variation of PCya production and biomass. Assuming a growth efficiency of 0.4, total protozoan production varied seasonally (189?C313?g?C?m^?2?day^?1) and was roughly half of particulate phytoplankton production. This study provides evidence that NF and PCya were tightly coupled in Lake Tanganyika and that herbivory by protists may be one of the reasons why this great lake has high productivity. Our results bring support to the idea that microbial herbivory is a major process in oligotrophic freshwater systems.     

Copepod grazing during an iron-induced diatom bloom in the Antarctic Circumpolar Current (EisenEx): I. Feeding patterns and grazing impact on prey populations

Feeding activity, selective grazing and the potential grazing impact of two dominant grazers of the Polar Frontal Zone, Calanus simillimus and Rhincalanus gigas, and of copepods < 2 mm were investigated with incubation experiments in the course of an iron fertilized diatom bloom in November 2000. All grazers were already actively feeding in the low chlorophyll waters prior to the onset of the bloom. C. simillimus maintained constant clearance rates and fed predominantly on diatoms. R. gigas and the small copepods strongly increased clearance and ingestion of diatoms in response to their enhanced availability. All grazers preyed on microzooplankton, most steadily on ciliates, confirming the view that pure herbivory appears to be the exception rather than the rule in copepod feeding. The grazers exhibited differences in feeding behavior based on selectivity indices. C. simillimus and R. gigas showed prey switching from dinoflagellates to diatoms in response to the phytoplankton bloom. All grazers most efficiently grazed on large diatoms leading to differences in daily losses for large and small species, e.g. Corethron sp. or Thalassionema nitzschioides. Species-specific diatom mortality rates due to grazing suggest that the high feeding activity of C. simillimus prior to and during the bloom played a role in shaping diatom population dynamics.     

Ciliate community structure, diversity and trophic role in offshore sediments from the Yellow Sea

We investigated the community structure, diversity and trophic role of ciliates in the sediments from 48 stations in the Yellow Sea using Ludox density centrifugation and quantitative protargol stain. The ciliate abundance ranged from 1 to 221cellscm(-3) and biomass from 0.0001 to 0.47μgCcm(-3) in the upper 8cm of the sediments. On average, 77% of ciliate abundance and 81% of biomass were distributed in the 0-2cm sediment layers, while the respective proportions were only about 6% and 3% in the 5-8-cm layers. Among the 198 morphospecies, Prostomatea was the most dominant group accounting for 45% of the total abundance and 58% of the total biomass. Carnivorous ciliates constituted the primary feeding type, occupying about 64% of the total biomass, followed by bacterivores (21%), algivores (12%) and omnivores (3%). The ciliate abundance and biomass in the upper 5cm of sediments were two orders of magnitude higher than those in the upper 10m of the Yellow Sea water column. The estimated ciliate bacterivory and herbivory indicate that ciliate ingestion had little direct influence on bacterial standing stock but possibly had an important impact on diatoms in the sediments from the Yellow Sea.     

Nitric oxide mediates oxylipin production and grazing defense in diatoms

Diatom blooms are important features of productive marine ecosystems and are known to support higher trophic levels. However, when stressed or wounded, diatoms can produce oxylipin molecules known to inhibit the reproduction and development of copepods and decrease microzooplankton growth rates. Using oxylipin chemical treatments, lipidomic analysis and functional genomic approaches, we provide evidence that nitric oxide (NO) and oxylipin signalling pathways in diatoms respond to protist grazers, resulting in increased defence fitness and survival. Exposure of the diatom Phaeodactylum tricornutum to the dinoflagellate Oxyrrhis marina resulted in NO production by P. tricornutum and pronounced change in its dissolved oxylipin profile. Experimentally elevating levels of NO also resulted in increased oxylipin production, and lower overall grazing rates. Furthermore, O. marina preferentially grazed on P. tricornutum prey with lower levels of NO, suggesting that this molecule and its effect on oxylipin pathways play a key role in prey selection. Exposure of O. marina grazing on P. tricornutum to exogenous oxylipins also decreased grazing rates, which is consistent with a grazing deterrence role for these molecules. These results suggest that NO and oxylipin production help to structure diatom communities, in part by modulating interactions with microzooplankton predators.     

The significance of inter- and intraspecific variation in bacterivorous and herbivorous protists

This paper reviews the emerging evidence on the significance of inter- and intraspecific variation in the feeding behaviour of aquatic protists. Small heterotrophic nanoflagellates (HNF) have been identified as the primary bacterial consumers in most aquatic environments. Recent research using novel techniques such as flow cytometry and high resolution video microscopy revealed that their feeding strategies and grazing rates are diverse. There is an important conceptual difference between uptake rates measured in short-term (min to h) experiments and grazing rates averaged over a longer-term (d). This is because the latter are strongly affected by digestion rates which are species-specific, i.e. the same bacterial prey may be digested differently by various grazers, and the same predator may selectively digest variable prey. Planktonic ciliates are the most important algal consumers in many lakes and marine systems. Large species-specific differences in their feeding behaviour and growth rates have been documented for closely related species. Intraspecific variation, which is, most likely, caused by varying clonal composition may be as important as interspecific variation. Finally, there is some evidence that the individual variability within a given population is generally large, both among bacterivorous HNF and among herbivorous ciliates. The consequences of this diversity becoming apparent at the levels of the species, population, clone and individual need to be considered by aquatic ecologists in their conceptual models. This revised version was published online in August 2006 with corrections to the Cover Date.     

Complex interactions and different possible pathways among functional components of the aquatic microbial world in Farasan Archipelago,Southern Red Sea,Saudi Arabia

This work aims to outline the dynamics of trophic links between the three main microbial components (bacteria, nanoflagellates, and ciliates) of the Farasan Archipelago in order to establish a baseline for future research in this area. The Farasan Archipelago lies along the southwestern coast of the Saudi Arabia, southern Red Sea between 16°20′–17°10′N and 41°30′–42°30′E and had been declared as marine and terrestrial reserve by the year 1996. Three different sites were chosen for this study, with each site visited bimonthly for 18 months from September 2016 to February 2018. Bacteria, nanoflagellates and ciliates were enumerated in order to explore the complex interactions between the main microbial categories in sea waters of the Farasan Archipelago. High abundances were recorded during the present study for bacteria (8.7 × 10⁶ bacteria ml⁻¹), nanoflagellates (3.7 × 10⁴ TNAN ml⁻¹) and ciliates (40.4 ciliates ml⁻¹). The paper discusses the various potential pathways controlling the complex interactions between these microbial groups in this part of the southern Red Sea. It is concluded that a linear trophic chain consisting of bacteria; heterotrophic nanoflagellates; filter feeding ciliates is a major route by which the production of bacteria is transferred to the higher consuming levels, thereby confirming the high importance of t bottom-up control (food supply), alongside top-down control (predation) in regulating bacterial abundances in the Farasan Archipelago. During the present investigation, each nanoflagellate ingested between 11 and 87 bacteria in one hour, while each ciliate consumed between 20 and 185 nanoflagellates every hour. These calculated grazing rates of protistan eukaryotes confirmed the role of heterotrophic nanoflagellates as the main consumers of bacteria, and the role of ciliates as the major control for the heterotrophic nanoflagellate population dynamics, and thus the top predators within the microbial plankton assemblage in the Farasan Archipelago.     

Diel feeding pattern and prey selection of mesozooplankton on microplankton community

Mesozooplankton play an important role in transporting energy from microphytoplankton and microzooplankton to higher trophic levels. However there were few studies on the diel feeding patterns and prey selectivity of mesozooplankton. We conducted feeding experiments of mesozooplankton in the East China Sea to determine their respective diel feeding patterns on diatoms, ciliates and dinoflagellates, and to assess the contribution of these prey items to mesozooplankton diet. The results showed higher mesozooplankton grazing rates on ciliates and dinoflagellates than on diatoms at the day time, and the opposite pattern at the night time. A significant prey selection was observed, in which mesozooplankton positively selected ciliates and dinoflagellates during day and diatoms at night. The different grazing reactions of mesozooplankton toward each prey item might be related to the mobility of the prey. In all, microzooplankton (ciliates and dinoflagellates) provided the majority of the mesozooplankton carbon ingestion, even at a station dominated by small pennate diatoms. In particular, dinoflagellates are an important prey of mesozooplankton in the East China Sea and their contribution to the diet of mesozooplankton is unproportionate to their abundance.