Seasonal microbial processes in a high-latitude fjord (Kongsfjorden, Svalbard): I. Heterotrophic bacteria, picoplankton and nanoflagellates

Temporal dynamics of the microbial food web in the Barents Sea and adjacent water masses in the European Arctic are to a large extent unknown. Seasonal variation in stocks and production rates of heterotrophic bacteria and phototrophic and heterotrophic picoplankton and nanoflagellates was investigated in the upper 50 m of the high-latitude Kongsfjorden, Svalbard, during six field campaigns between March and December 2006. Heterotrophic bacteria, picoplankton and nanoflagellates contributed to ecosystem structure and function in all seasons. Activity within the microbial food web peaked during spring bloom in April, parallel to low abundances of mesozooplankton. In the nutrient-limited post-bloom scenario, an efficient microbial loop, fuelled by dissolved organic carbon from abundant mesozooplankton feeding on phytoplankton and protozooplankton, facilitated maximum integrated primary production rates. A tight microbial food web consisting of heterotrophic bacteria and phototrophic and heterotrophic picoplankton and nanoflagellates was found in the stratified water masses encountered in July and September. Microbial stocks and rates were low but persistent under winter conditions. Seasonal comparisons of microbial biomass and production revealed that structure and function of the microbial food web were fundamentally different during the spring bloom when compared with other seasons. While the microbial food web was in a regenerative mode most of the time, during the spring bloom, a microbial transfer mode represented a trophic link for organic carbon in time and space. The microbial food web’s ability to fill different functional roles in periods dominated by new and regenerated production may enhance the ecological flexibility of pelagic ecosystems in the present era of climate change.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Trophic coupling of the microbial and the classical food web in Lake Baikal,Siberia

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Structure and functioning of the microbial loop in a boreal reservoir

The abundance and biomass of the main components of the microbial plankton food web (“microbial loop”)—heterotrophic bacteria, phototrophic picoplankton and nanoplankton, heterotrophic nanoflagellates, ciliates and viruses, production of phytoplankton and bacterioplankton, bacterivory of nanoflagellates, bacterial lysis by viruses, and the species composition of protists—have been determined in summer time in the Sheksna Reservoir (the Upper Volga basin). A total of 34 species of heterotrophic nanoflagellates from 15 taxa and 15 species of ciliates from 4 classes are identified. In different parts of the reservoir, the biomass of the microbial community varies from 26.2 to 64.3% (on average 45.5%) of the total plankton biomass. Heterotrophic bacteria are the main component of the microbial community, averaging 63.9% of the total microbial biomass. They are the second (after the phytoplankton) component of the plankton and contribute on average 28.6% to the plankton biomass. The high ratio of the production of heterotrophic bacteria to the production of phytoplankton indicates the important role of bacteria, which transfer carbon of allochthonous dissolved organic substances to a food web of the reservoir.     

On the Importance of Planktonic Protozoans in the Eutrophication Process of the Baltic Sea

Analysing the results of various authors recent studies in the pelagic region of the Baltic revealed that protozoan biomass is in the same range or even higher than metazooplankton biomass. The dominant groups of planktonic protozoans are heterotrophic pico- and nanoflagellates (various taxonomic groups), large heterotrophic flagellates (mainly dinoflagellates) and ciliates. Regularly the spring bloom of phytoplankton is accompanied by a maximum of protozoan biomass which declines in early summer as a result of intensive grazing pressure by metazooplankton and changing food conditions. The analysis of results from different stations indicated that biomasses of protozoans increase with an increasing degree of eutrophication. Several trophic levels within the microbial web should be added to the traditional view on the pelagic food web of the Baltic. Our knowledge regarding the quantitative aspect of the microbial matter flux of the Baltic is very limited up to now and complex ecological (and taxonomical) studies using standardized methods including all protozoan components are necessary. Protozoans (various trophic groups and levels), besides bacteria, should be viewed as the metabolically most active heterotrophic component in the pelagic region of the Baltic, their activity should increase with an increasing degree of eutrophication.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Isotopic analysis of the sources of organic carbon for zooplankton in shallow subarctic and arctic waters

Shallow high-latitude lakes and ponds are usually characterized by an oligotrophic water column overlying a biomass-rich, highly productive benthos. Their pelagic food webs often contain abundant zooplankton but the importance of benthic organic carbon versus seston as their food sources has been little explored. Our objectives were to measure the δ¹³C and δ¹⁵N isotopic signatures of pelagic and benthic particulate organic matter (POM) in shallow water bodies in northern Canada and to determine the relative transfer of this material to zooplankton and other aquatic invertebrates. Fluorescence analysis of colored dissolved organic matter (CDOM) indicated a relatively strong terrestrial carbon influence in five subarctic waterbodies whereas the CDOM in five arctic water columns contained mostly organic carbon of autochthonous origin. The isotopic signatures of planktonic POM and cohesive benthic microbial mats were distinctly different at all study sites, while non-cohesive microbial mats often overlapped in their δ¹³C signals with the planktonic POM. Zooplankton isotopic signatures indicated a potential trophic link with different fractions of planktonic POM and the non-cohesive mats whereas the cohesive mats did not appear to be used as a major carbon source. The zooplankton signals differed among species, indicating selective use of resources and niche partitioning. Most zooplankton had δ¹³C values that were intermediate between the values of putative food sources and that likely reflected selective feeding on components of the pelagic or benthic POM. The results emphasize the likely importance of benthic-pelagic coupling in tundra ecosystems, including for species that are traditionally considered pelagic and previously thought to be dependent only on phytoplankton as their food source.     

Phytoplankton food quality control of planktonic food web processes

We developed a mechanistic model of nutrient, phytoplankton, zooplankton and fish interactions to test the effects of phytoplankton food quality for herbivorous zooplankton on planktonic food web processes. When phytoplankton food quality is high strong trophic cascades suppress phytoplankton biomass, the zooplankton can withstand intense zooplanktivory, and energy is efficiently transferred through the food web sustaining higher trophic level production. Low food quality results in trophic decoupling at the plant-animal interface, with phytoplankton biomass determined primarily by nutrient availability, zooplankton easily eliminated by fish predation, and poor energy transfer through the food web. At a given nutrient availability, food quality and zooplanktivory interact to determine zooplankton biomass which in turn determines algal biomass. High food quality resulted in intense zooplankton grazing which favored fast-growing phytoplankton taxa, whereas fish predation favored slow-growing phytoplankton. These results suggest algal food quality for herbivorous zooplankton can strongly influence the nature of aquatic food web dynamics, and can have profound effects on water quality and fisheries production. Handling editor: D. Hamilton     

Effects of Additions of DOC on Pelagic Biota in a Clearwater System: Results from a Whole Lake Experiment in Northern Sweden

An oligotrophic clearwater lake, initially characterized by a pronounced dominance of autotrophic phytoplankton and mostly by one species, the green alga Botryococcus, was subject to additions of dissolved organic carbon in the form of white sugar (sucrose) during two consecutive years. The hypothesis tested was that it is organic carbon per se, and not other possible effects of humic substances, that determines the differences in structure of the planktonic ecosystem between humic and clearwater lakes. The additions of DOC resulted in a significant increase in bacterial biomass and a decrease in the biomass of autotrophic phytoplankton. The biomass of mixotrophic and heterotrophic flagellates instead increased significantly, whereas no effects were found to propagate to higher trophic levels. As a result of the changes among biota, total planktonic biomass also decreased to a level typical of nearby humic lakes. We suggest that it is the carbon component of humic material and its utilization by bacterioplankton that determines the structure and function of the pelagic food web in humic lakes.     

Benthic and pelagic food resources for zooplankton in shallow high-latitude lakes and ponds

1. Shallow lakes and ponds are a major component of the northern landscape and often contain a high zooplankton biomass despite clear waters that are poor in phytoplankton. 2. In this study we quantified zooplankton food sources and feeding rates in the shallow waters of two contrasting high‐latitude biomes: subarctic forest tundra (Kuujjuarapik, Quebec) and high arctic polar desert (Resolute, Nunavut). Five substrate types were tested (beads, bacteria, picophytoplankton, filamentous plankton and microbial mats). Special attention was given to the role of benthos, a component that is usually poorly integrated into models of aquatic foodwebs. 3. Consistent with observations elsewhere in the circumpolar region, high concentrations of adult macrozooplankton occurred in all sites (up to 17 100 crustaceans m^?3) while phytoplankton concentrations and primary productivity were low. The communities were composed of multiple species, including Daphnia middendorfiana, Hesperodiaptomus arcticus, Leptodiaptomus minutus, Artemiopsis stefanssoni and Branchinecta paludosa. 4. Detritus made 89–98% of the planktonic resource pool and bacteria contributed the highest biomass (up to 29 mg C m^?3) of the planktonic food particles available to zooplankton. Benthic resources were dominated by microbial mats that grew in nutrient‐rich conditions at the base of the ponds and which dominated overall ecosystem biomass and productivity. 5. All species were flexible in their feeding but there were large, order of magnitude differences in clearance rates among taxa. These differences likely resulted from different grazing strategies among cladocerans, copepods and fairy shrimps, and possibly also from adaptation to specific food types and size ranges that occur locally in these waters. 6. The subarctic cladocerans Ceriodaphnia quadrangula and D. middendorfiana, and the arctic fairy shrimp B. paludosa were observed to graze directly on the microbial mats and the feeding experiments confirmed their assimilation of benthic substrates. The other zooplankton species showed a more pelagic feeding mode but were capable of using microbial mat filaments, thus may be indirectly linked to benthic processes via resuspension. 7. Our study indicates that the classical aquatic food web in which phytoplankton provide the sole production base for grazers does not apply to northern shallow lakes and ponds. Instead, microbial mats increase the physical complexity of these high latitude ecosystems and likely play a role in sustaining their high zooplankton biomass.     

The microbial loop in the planktonic communities in lakes with various trophic status

The structure of planktic trophic chains was studied in eight lakes of European Russia and five lakes in Central Asia. The lakes differed in the level of productivity, morphometric parameters, and the type of agitation and mineralization. It is found that the microbial loop of picophototrophic organisms, bacteria, heterotrophic flagellates, infusoria, and viruses constitutes 12.3-64.7% of the total plankton biomass. Positive correlation between the biomass of microbial community and the primary production of phytoplankton is observed, whereas no relation is revealed between the share of microorganisms in the plankton biomass and the trophic status of the water body. The presence of a great number of cladocerans decreased the role of the microbial loop in the structural organization of the planktic community. Heterotrophic flagellates consuming 3-81% of daily bacterial production were the principal cause of bacteria elimination only in some of the studied water bodies.     

Unimodal patterns of microbial communities with eutrophication in Mediterranean shallow lakes

The role of the microbial communities in the classical planktonic food web and its response to eutrophication in shallow lakes is still contradictory. Mediterranean shallow lakes with different eutrophication levels were sampled to study the influence of eutrophication on the microbial food web (MFW) and their contribution to the planktonic food web. Percentage of ciliate biomass in the metazooplankton (MZP) showed a U-shaped trend with eutrophication, with maximum at both ends of the chlorophyll-a (Chla) gradient. The MZP to phytoplankton ratio demonstrated a unimodal pattern with minimum values at the two ends of the Chla gradient and maximum values in the Chla range 5-10 μg l^?1. In contrast, the MFW to phytoplankton ratio reached its minimum in the central part of the Chla gradient and maximum values at the extremes of the gradient. These patterns support the hypothesis that the relative importance of bacteria and ciliates is lowest in mesotrophic shallow lakes, and highest in oligotrophic and hypereutrophic systems. These results stress the importance of protozoan in the trophic web, and indicate it is essential to include this group, especially ciliates, when quantifying zooplankton in warm shallow lakes.     

Testing predictions of the lake food web theory on pelagic communities of Australian reservoirs

Several predictions of the theory developed for pelagic food webs of the Northern Hemisphere were tested on water bodies of Eastern Australia. Eleven reservoirs, representing trophic and latitudinal gradients were sampled for nutrients, phytoplankton, zooplankton and pelagic fish. Two models of regression analysis, which analysed possible interactions between trophic levels were based on different sets of data. In one, each reservoir was represented by only one pair of observations – annual mean or single observation (“regional model”). In the other, seasonal means of four frequently sampled reservoirs similar in productivity were used (“temporal model”). Significant variation in total phytoplankton biovolume (TPB) was predicted by total phosphorus concentration (TP), total nitrogen concentration (TN), mean crustacean length and acoustic biomass of planktivorous fish in both models. This suggested that nutrient limitation, zooplankton grazing and positive effects of fish were probably important in controlling the biomass of primary producers at both regional and temporal scales. In the regional model, the biomass of fish was also negatively correlated with Daphnia biomass and mean crustacean length, suggesting that the trophic cascade hypothesis may be applicable to Eastern Australia for the considered range of reservoir productivities. The biovolume of cyanobacteria was not correlated to any variables tested in the regional model. In contrast, nutrient and food web variables had significant effects on cyanobacterial biovolume in the temporal model. This suggested that factors governing seasonal succession were probably more important for cyanobacteria than variation in reservoir productivity or location. Contrary to previous views, no negative relationship between total biomass of zooplankton and TPB was found in both models, suggesting that the community structure of zooplankton rather than its total biomass mediates top‐down effects. Many predictions of the food web theory remained robust in spite of substantial differences in animal taxonomy and physical environment of Australian ecosystems.     

OPINION Manipulating lake community structure: where do we go from here?

SUMMARY. 1 More than 10 years experience with whole lake pelagic manipulation has suggested some general trends applicable to all freshwater pelagic communities and some specific trends related to lake depth.
2 Among the general trends is the observation that the trophic cascade is strongly damped. This means that changes in phytoplankton biomass can be assured only when the fish community is strongly manipulated.
3 Among the depth related trends is the observation that in shallow lakes, changes in fish community structure are more likely to have cascading impacts on phytoplankton than are changes in deep lakes.
4 In shallow lakes, fish removal frequently results in decreased turbidity which is associated with the development of dense macrophyte populations and significant reductions of algal standing stocks. The mechanisms involve: increased grazing by zooplankton, the removal of fish induced bioturbation and nutrient recycling, and direct and indirect macrophyte effects (shading, zooplankton refuges and competition for nutrients).
5 In shallow lakes, where planktivore biomass can be regulated and macrophyte development is acceptable, fish biomanipulalions are likely to result in reduced algal populations and improved water quality.
6 In deep lakes, where macrophytes are not as important, long-term effects of fish manipulations are strongly dependent upon the probability of non-grazable algal bloom development. This is determined by many factors (chemical, physical and grazer related) which modify the impact that grazers have on phytoplankton biomass.
7 In deep lakes, successful fish biomanipulations may only be effective when chemical and physical factors are altered to produce algal species compositions that permit strong top-down control of prey by predators.     

Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem: Insight from carbon and nitrogen stable isotopes

Stable carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) are used to study the trophic structure of food web in the Yellow Sea and East China Sea ecosystem. The trophic continuum of pelagic food web from phytoplankton to top preyer was elementarily established, and a trophic structure diagram in the Yellow Sea and East China Sea was outlined in combination with carbon isotopic data of benthic organisms, which is basically consistent with and makes some improvements on the simplified Yellow Sea food web and the trophic structure diagram drawn based on the biomass of main resource population during 1985–1986. This result indicates that the stable isotope method is a potential useful means for further studying the complete marine food web trophic continuum from viruses to top predators and food web stability.     

Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem: Insight from carbon and nitrogen stable isotopes

The determination of trophic level for the biology in a marine ecosystem is very important as alteration of its structure and function may be reflected in the tro-phic level of component species. A change in trophic level indicates variation in an organism’s feeding bi-ology or in the pathway of energy flow from primary producers to the consumer. The gut content analysis is a traditional method for studying trophodynamics of food web in marine ecosystems. Species composition and amounts in al…     

Plankton biomass partitioning in a eutrophic subtropical lake: comparison with results from temperate lake ecosystems

Plankton were sampled for 6 years in a subtropical eutrophiclake in FL, USA, and absolute and relative carbon biomass wasdetermined for bacteria, phytoplankton, heterotrophic and phototrophicnanoflagellates, ciliates, rotifers and crustacean zooplankton.We compared the results with findings from a comprehensive studyof carbon biomass partitioning in eutrophic German lakes withelucidate common patterns and differences. Similarities betweenthe temperate and subtropical systems included: similar seasonaldynamics, with maximal carbon biomass of nanoflagellates andmetazoan zooplankton in spring and phytoplankton in summer toautumn, yearly averaged carbon occurring mainly in the phytoplanktonand phytoplankton accounting for a much greater proportion ofcarbon than bacteria. There also were differences: the Floridalake had lower absolute and relative carbon biomass in crustaceanzooplankton, stronger dominance of protozoa in total grazercarbon biomass, a lower ratio of zooplankton to phytoplanktoncarbon and almost a monoculture of predation-resistant copepods(versus a relatively balanced distribution of carbon among cladocerans,copepods and rotifers in the temperate lakes). The subtropicallake also had 4-fold higher relative biomass of small filamentouscyanobacteria in its phytoplankton, which we attribute to lightlimitation. Although the Florida and German studies did notmeasure biomass of planktivorous fish, the differences observedhere are consistent with a recent hypothesis that fish predationexerts stronger top–down control on the pelagic food webin subtropical lakes than in temperate lakes of similar trophicstatus.     

Algal picoplankton production and contribution to food-webs in oligotrophic British Columbia lakes

The pelagic communities of two contrasting oligotrophic lakes in British Columbia were studied to determine why an interior, dimictic lake (Quesnel) supports a greater biomass of zooplankton and produces larger planktivorous sockeye salmon (Oncorhynchus nerka) than a coastal warm-monomictic lake (Sproat). The ultra-oligotrophic status and differing planktivore densities in Sproat Lake increased the relative importance of algal picoplankton, diminished the abundance of large zooplankton, and increased the significance of rotifers and other small-bodied zooplankton. These picoplankton based food webs result in longer, indirect and less efficient pathways of carbon flow from phytoplankton to fish. In contrast, Quesnel Lake is a more productive oligotrophic lake and its pelagic food webs are based more on nanoplankton and small microphytoplankton that support larger-bodied zooplankton (Daphnia, Diaptomus), and a more direct and efficient two-step transfer to fish. The greater variability of the annual recruitment of sockeye fry in interior lakes may keep zooplankton communities in a non-steady state, this in turn may perpetuate the occurrence of quadrennial cyclic dominance in adult salmon returning to these systems.     

Mixing, stratification and the fate of primary production in an oligotrophic multibasin lake system (Quebec, Canada)

Impacts of mixing and stratification on the fate of primaryproduction were studied in an oligotrophic lake by comparingthe size-distributions of phytoplankton standing stock and productionin two basins, only one of which experiences seasonal thermalstratification. In both basins, the phytoplankton was dominatedby small cells (pico- and nanoplankton). The contribution ofpicoplankton to both biomass and production remained relativelyconstant throughout the season in both basins. Seasonal variationsin the size structure of phytoplankton communities do not agreewith the paradigm of dominance by small cells during summerstratification and dominance of larger cells during spring andfall mixing events. Nutrient control of productivity throughmixing and stratification is unlikely to affect the structureof phytoplankton communities when nutrients (allochthonous)derived from the catchment basin or sediments are in short supply.In such environments, nutrients (autochthonous) are largelyderived in the lake through heterotrophic food web processessuch as grazing, excretion and decomposition. Maximum ratesof production and losses in July and August in both basins areconsistent with increased regeneration and may represent a responseof larger-sized cells to higher nutrient availability resultingfrom enhanced grazing on picoplankton. The high correlationbetween the rates of loss and of potential growth for the phytoplanktoncommunity during all sampling periods, and the relative constancyof the picoplankton biomass, leads us to propose a long-term,steady-state equilibrium in the phytoplankton community underthe control of grazing by herbivores and/or other loss processes.     

Influences of fish on food web structure and function in mountain lakes

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