Size‐based interactions across trophic levels in food webs of shallow Mediterranean lakes

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Live labeling technique reveals contrasting role of crustacean predation on microbial loop in two large shallow lakes

We tested the hypotheses that the ciliate assemblages in moderately eutrophic lake are controlled by the effective crustacean predation, and the high abundances of planktonic ciliates in highly eutrophic and turbid lake are due to insufficient regulation by crustacean zooplankton. A food tracer method coupled with natural assemblage of microciliates labeled with fluorescent microparticles was used to measure the cladoceran and copepod predation rates on planktonic ciliates and to estimate the carbon flow between the ciliate–crustacean trophic links. The results revealed that the microciliates (15–40 μm) were consumed by all dominant cladoceran and copepod species in both the lakes studied, mainly by Chydorus sphaericus and cyclopoid copepods in Lake Võrtsjärv, and by Daphnia spp. and Bosmina spp. in Lake Peipsi. The grazing loss in moderately eutrophic Peipsi indicated strong top-down control of ciliates mainly by cladocerans. The extraordinary abundant population of planktonic ciliates having a predominant role in the food web in highly eutrophic and turbid Võrtsjärv is explained by the measured low crustacean predation rates on ciliates. The estimated carbon flow from the ciliates to crustaceans suggest that in eutrophic lakes majority of the organic matter channeled via metazooplankton to higher trophic levels may originate from the microbial loop.     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

Significance of predation by protists in aquatic microbial food webs

Predation in aquatic microbial food webs is dominated by phagotrophic protists, yet these microorganisms are still understudied compared to bacteria and phytoplankton. In pelagic ecosystems, predaceous protists are ubiquitous, range in size from 2 μm flagellates to >100 μm ciliates and dinoflagellates, and exhibit a wide array of feeding strategies. Their trophic states run the gamut from strictly phagotrophic, to mixotrophic: partly autotrophic and partly phagotrophic, to primarily autotrophic but capable of phagotrophy. Protists are a major source of mortality for both heterotrophic and autotrophic bacteria. They compete with herbivorous meso- and macro-zooplankton for all size classes of phytoplankton. Protist grazing may affect the rate of organic sinking flux from the euphotic zone. Protist excretions are an important source of remineralized nutrients, and of colloidal and dissolved trace metals such as iron, in aquatic systems. Work on predation by protists is being facilitated by methodological advances, e.g., molecular genetic analysis of protistan diversity and application of flow cytometry to study population growth and feeding rates. Examples of new research areas are studies of impact of protistan predation on the community structure of prey assemblages and of chemical communication between predator and prey in microbial food webs. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Fish communities and food web interactions in some shallow Mediterranean lakes

Hydrobiologia - Fish communities of five shallow Spanish Mediterranean lakes were studied and stomach content analysed to determine the foraging pattern of each species and the influence on the...     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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Combined stable isotope and gut contents analysis of food webs in plant-dominated, shallow lakes

Summary 1. To determine feeding links between primary producers, invertebrates and fish, stable isotope analyses and gut content analyses of fish were conducted on the components of four shallow, eutrophic to hypertrophic, plant-dominated lakes.
2. Although separation of basal resources was possible, the diets of both fish and invertebrates were broad, comprising food from different compartments (planktonic, epiphytic/benthic), as well as from different trophic levels.
3. Mixing models were used to determine the extent to which periphyton production supported higher trophic levels. Only one species of invertebrate relied upon periphyton production exclusively.
4. Fish density affected the diets of invertebrates. The response was different for planktonic and epiphytic/benthic invertebrates. The proportion of periphyton production in the diets of zooplankton appeared to increase with fish density, whilst it decreased for other invertebrates.
5. As all zooplankton samples were collected in the open water at dusk, these results are further evidence for the diurnal horizontal migration of zooplankton. Although not conclusive, they are consistent with a behavioural response by invertebrates and zooplankton in the presence of fish.     

Energetics of microbial food webs

The energetic demand of microorganisms in natural waters and the flux of energy between microorganisms and metazoans has been evaluated by empirical measurements in nature, in microcosms and mesocosms, and by simulation models. Microorganisms in temperate and tropical waters often use half or more of the energy fixed by photosynthesis. Most simulations and some experimental results suggest significant energy transfer to metazoans, but empirical evidence is mixed. Considerations of the range of growth yields of microorganisms and the number of trophic transfers among them indicate major energy losses within microbial food webs. Our ability to verify and quantify these processes is limited by the variability of assimilation efficiency and uncertainty about the structure of microbial food webs. However, even a two-step microbial chain is a major energy sink. As an energetic link to metazoans, the detritus food web is inefficient, and its significance may have been overstated. There is not enough bacterial biomass associated with detritus to support metazoan detritivores. Much detritus is digestible by metazoans directly. Thus, metazoans and bacteria may to a considerable degree compete for a common resource. Microorganisms, together with metazoans, are important to the stability of planktonic communities through their roles as rapid mineralizers of organic matter, releasing inorganic nutrients. The competition for organic matter and the resultant rapid mineralization help maintain stable populations of phytoplankton in the absence of advective nutrient supply. At temperatures near O °C, bacterial metabolism is suppressed more than is the rate of photosynthesis. As a result, the products of the spring phytoplankton bloom in high-temperate latitudes are not utilized rapidly by bacteria. At temperatures below 0°C microbial food webs are neither energy sinks or links: they are suppressed. Because the underlying mechanism of low-temperature inhibition is not known, we cannot yet generalize about this as a control of food web processes. Microorganisms may operate on several trophic levels simultaneously. Therefore, the realism of the trophic level concept and the reality of the use of ecological efficiency calculations in ecosystem models is questionable.     

Phagotrophic phytoflagellates in microbial food webs

Phagotrophy by pigmented flagellates is known from the literature but has recently been rediscovered in the context of microbial food webs. Particle ingestion rates were found to be equivalent for pigmented and nonpigmented microflagellates in both field and laboratory studies. Ingestion rates of the chrysophytes Ochromonas danica, O. minuta, and Poterioochromonas malhamensis, the dinoflagellate Peridinium inconspicuum, and the cryptophytes Cryptomonas ovata and C. erosa were compared with those of two nonpigmented Monas species using 0.57 μm polystyrene beads as a food source. Ingestion rates were 0.31 to 3.17 beads/cell/h and filtration rates were 10⁻⁷ to 10⁻⁸ ml/cell/h with no detectable difference between pigmented and nonpigmented forms. Ingestion rates in unpigmented Monas species showed a linear increase with increasing particle concentration from 1.9 × 10⁶ to 1.6 × 10⁷ beads/ml. Light and DOC levels in the range of those encountered by phytoflagellates in the field also influenced laboratory measurements of bead ingestion by Poterioochromonas malhamensis. Ingestion rates decreased and photosynthesis increased over the natural PAR light range from 0 to 1800 microeinsteins/s/m². At 40 microeinsteins/s/m² maximum ingestion rates and high rates of photosynthesis occurred simultaneously. Ingestion rates decreased above 4 mgC/l supplied as glucose. DOC levels commonly occurring in Lake Oglethorpe range from 3.5 to 10.0 mgC/l. These studies suggest that mixotrophy, the trophic utilization of particulate food and dissolved organic matter as well as photosynthetically fixed organic matter, is a balanced process that can be regulated by environmental conditions. In field studies during a chrysophyte bloom, phytoflagellate grazing exceeded heterotrophic microflagellate grazing and constituted up to 55% of the bactivory of all microflagellates, ciliates, rotifers, and crustaceans combined. Neither bacterial abundance, light nor temperature were good predicters of grazing rates for the phagotrophic phytoflagellate association as a whole during this unstratified period. Phagotrophs are often most abundant at the metalimnetic plate during stratification.     

Responses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment

1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte‐dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L^?1 P and 10 mg L^?1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within‐treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD < 50 μm). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes.     

Salinity effects on macroinvertebrate assemblages and waterbird food webs in shallow lakes of the Wyoming High Plains

We determined the biomass and community structure of macroinvertebrates (>500 µm) associated with macrophytes, sediments, and unvegetated open water in three oligosaline (0.8 to 8.0 mS cm^–1) and three mesosaline (8.0 to 30.0 mS cm^–1) lakes in the Wyoming High Plains, USA. Total biomass of epiphytic and benthic invertebrates did not change with salinity, but biomass of macroinvertebrate zooplankton in open water was significantly higher in mesosaline lakes. Community composition of invertebrates differed between the two salinity categories: large grazer/detritivores (gastropods and amphipods) were dominant in oligosaline lakes, whereas small planktivores and their insect predators were more prevalent in mesosaline lakes. Both direct physiological effects of salinity, as well as a shift in the form of primary production from macrophytes to phytoplankton, probably explain these changes in community composition. Salinity effects on invertebrate communities appear to be less important to top avian consumers than are costs of osmoregulation.     

Comparison of pelagic food webs in lakes along a trophic gradient and with seasonal aspects: influence of resource and predation

Composition and seasonal dynamics of phytoplankton, bacteria,and zooplankton (including heterotrophic flagellates, ciliates,rotifers and crustaceans) were studied in 55 lakes in NorthernGermany with different trophic status, ranging from mesotrophicto hypertrophic. Mean abundance and biomass of all groups increasedsignificantly with trophic level of the lake, but bacteria andmetazooplankton showed only a weak correlation and a slightincrease with chlorophyll concentration. Composition of phytoplanktonshowed a dominance of cyanobacteria in hypertrophic lakes, whereasthe importance of chrysophytes and dinophytes decreased withan increase in trophic status. Protozoans (heterotrophic flagellatesand ciliates) made up 24% (mesotrophic lakes) to 42% (hypertrophiclakes) of total zooplankton biomass on average, and were dominatedby ciliates (62–80% of protozoan biomass). Seasonally,protozoans can build up to 60% of zooplankton biomass in spring,when heterotrophic flagellates can contribute     

Modelling food webs and nutrient cycling in agro-ecosystems

Agricultural practices affect the spatial patterns and dynamics of the decomposition of soil organic matter and the availability of plant-limiting nutrients. The biological processes underlying these patterns and dynamics are the trophic interactions among the organisms in the soil community food web. Food web models simulate nutrient flow rates close to observed rates and clarify the role of the various groups of organisms in the cycling of nutrients. Several large interdisciplinary programs are currently focusing on these interactions, with a view to developing and managing sustainable forms of agriculture.     

Aggregation and disaggregation of microbial food webs

Models of the microbial food web have generally used compartments aggregated by general body size and gross taxonomy. It has been assumed that these also reflect guilds or holons. Generally, results of simulation or analysis based on this structure have been reasonably well validated. Herein I summarize why the aggregations may be justified and what may be learned from disaggregation.     

Structure, biomass, production and depth distribution of periphyton on artificial substratum in shallow lakes with contrasting nutrient concentrations

1. To examine how the vertical distribution of periphytic biomass and primary production in the upper 0–1 m of the water column changes along an inter‐lake eutrophication gradient, artificial substrata (plastic strips) were introduced into the littoral zones of 13 lakes covering a total phosphorus (TP) summer mean range from 11 to 536 μg L^?1. Periphyton was measured in July (after 8 weeks) and September (after 15 weeks) at three water depths (0.1, 0.5 and 0.9 m). 2. Periphyton chlorophyll a concentration and dry weight generally increased with time and the communities became more heterotrophic. Mean periphytic biomass was unimodally related to TP, reaching a peak between 60 and 200 μg L^?1. 3. The proportion of diatoms in the periphyton decreased from July to September. A taxonomic shift occurred from dominance (by biovolume) of diatoms and cyanobacteria at low TP to dominance of chlorophytes at intermediate TP and of diatoms (Epithemia sp.) in the two most TP‐rich lakes. 4. The grazer community in most lakes was dominated by chironomid larvae and the total biomass of grazers increased with periphyton biomass. 5. Community respiration (R), maximum light‐saturated photosynthetic rate (P_max), primary production and the biomass of macrograzers associated with periphyton were more closely related to periphyton biomass than to TP. Biomass‐specific rates of R, P_max and production declined with increasing biomass. 6. Mean net periphyton production (24 h) was positive in most lakes in July and negative in all lakes in September. Net production was not related to the TP gradient in July, but decreased in September with increasing TP. 7. The results indicate that nutrient concentrations alone are poor predictors of the standing biomass and production of periphyton in shallow lakes. However, because periphyton biomass reaches a peak in the range of phosphorus concentration in which alternative states occur in shallow lakes, recolonisation by submerged macrophytes after nutrient reduction may potentially be suppressed by periphyton growth.     

Trends in research on shallow water food webs

Making sense of the spider–web networks of interactions between species in food webs has been a major pre-occupation of ecologists over the last 60 years. This review describes the early attempts to reduce this complexity through the grouping of individual taxa into functional categories (such as trophic levels), through adopting the energy flow or systems approach as epitomised by the International Biological Programme, and most recently by the derivation of web statistics by food web theorists. The strengths and weaknesses of these approaches are discussed in relation to empirical field experiments for unravelling the processes responsible for organising communities and an assessment made of the representation of these approaches in the marine biological literature.     

Trophic levels in community food webs

Summary Four concepts are considered for the trophic level of a species in a community food web. The long-way-up-level (or LU-level) of species A is the length of the longest simple food chain from a basal species (one with no prey in the web) to A. (A simple chain is a chain that does not pass through any given species more than once.) The short-way-up-level (SU-level) of species A is the length of the shortest chain from a basal species to A. The long-way-down-level (LD-level) of species A is the length of the longest simple chain from species A to a top species (one with no consumers in the web). The short-way-down-level (SD-level) of species A is the length of the shortest chain from species A to a top species. The stratigraphy of a web is the analogue for species of the pyramid of numbers for individuals: it is the frequency distribution of species according to level. The LU-, SU-, LD-, and SD-stratigraphies of the seven webs in the Briand-Cohen collection with 30 or more trophic species reveal no species with LU-level or LD-level more than 6, no species with SU-level more than 3, and no species with SD-level more than 2. In all seven webs, SD-levels are stochastically less than SU-levels: species tend to be closer to a top predator than to a basal species. Two stochastic models of food web structure (the cascade model and the homogeneous superlinear model) correctly predict that 95% or more of species should have LU-level and LD-level in the range 0–6. The models also correctly predict some details of the distribution of species in the SU- and SD-stratigraphies, particularly the fraction of species in level 1. The models do not, in general, correctly predict the distribution of species within the range 0–6 of LU-levels and LD-levels.     

Drought-induced changes in nutrient concentrations and retention in two shallow Mediterranean lakes subjected to different degrees of management

While extensive knowledge exists on the relationship between nutrient loading and nutrient concentrations in lakes in the cold temperate region, few studies have been conducted in warm lakes, not least in warm arid lakes. This is unfortunate as a larger proportion of the world’s lakes will be situated in arid climates in the future due to climate change and a larger proportion will suffer from a higher frequency of intensive drought. We conducted a comprehensive 11–13 year mass balance study in two interconnected shallow Mediterranean lakes in Turkey, covering a period with substantial changes in climate conditions. The upstream lake was only affected by natural changes in nutrient loading, while the downstream lake was additionally influenced by sewage diversion and restoration by fish removal. Contrasting to experience from north temperate lakes we found an increase in in-lake concentrations of total phosphorus and inorganic nitrogen (ammonia as well as nitrate) in dry years despite lower external nutrient loading, and submerged macrophytes did not increase the nitrogen retention capacity of the lakes. In contrast, fish removal modulated the nitrogen concentration as in north temperate lakes, but the effect was not long-lasting. Our results suggest that climate warming reduces the nutrient retention capacity of shallow lakes in the Mediterranean and exacerbates eutrophication. Lower thresholds of nutrient loading for shifting turbid shallow lakes to a clear water state are therefore to be expected in arid zones in a future warmer climate, with important management implications.