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.     

Food web architecture and basal resources interact to determine biomass and stoichiometric cascades along a benthic food web

Understanding the effects of predators and resources on primary producers has been a major focus of interest in ecology. Within this context, the trophic cascade concept especially concerning the pelagic zone of lakes has been the focus of the majority of these studies. However, littoral food webs could be especially interesting because base trophic levels may be strongly regulated by consumers and prone to be light limited. In this study, the availability of nutrients and light and the presence of an omnivorous fish (Hyphessobrycon bifasciatus) were manipulated in enclosures placed in a humic coastal lagoon (Cabiúnas Lagoon, Macaé - RJ) to evaluate the individual and interactive effects of resource availability (nutrients and light) and food web configuration on the biomass and stoichiometry of periphyton and benthic grazers. Our findings suggest that light and nutrients interact to determine periphyton biomass and stoichiometry, which propagates to the consumer level. We observed a positive effect of the availability of nutrients on periphytic biomass and grazers' biomass, as well as a reduction of periphytic C∶N∶P ratios and an increase of grazers' N and P content. Low light availability constrained the propagation of nutrient effects on periphyton biomass and induced higher periphytic C∶N∶P ratios. The effects of fish presence strongly interacted with resource availability. In general, a positive effect of fish presence was observed for the total biomass of periphyton and grazer's biomass, especially with high resource availability, but the opposite was found for periphytic autotrophic biomass. Fish also had a significant effect on periphyton stoichiometry, but no effect was observed on grazers' stoichiometric ratios. In summary, we observed that the indirect effect of fish predation on periphyton biomass might be dependent on multiple resources and periphyton nutrient stoichiometric variation can affect consumers' stoichiometry.     

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.     

Indirect and direct controls of macroinvertebrates and small fish by abiotic factors and trophic interactions in the Florida Everglades

1. The roles of nutrients, disturbance and predation in regulating consumer densities have long been of interest, but their indirect effects have rarely been quantified in wetland ecosystems. The Florida Everglades contains gradients of hydrological disturbance (marsh drying) and nutrient enrichment (phosphorus), often correlated with densities of macroinvertebrate infauna (macroinvertebrates inhabiting periphyton), small fish and larger invertebrates, such as snails, grass shrimp, insects and crayfish. However, most causal relationships have yet to be quantified. 2. We sampled periphyton (content and community structure) and consumer (small omnivores, carnivores and herbivores, and infaunal macroinvertebrates inhabiting periphyton) density at 28 sites spanning a range of hydrological and nutrient conditions and compared our data to seven a priori structural equation models. 3. The best model included bottom‐up and top‐down effects among trophic groups and supported top‐down control of infauna by omnivores and predators that cascaded to periphyton biomass. The next best model included bottom‐up paths only and allowed direct effects of periphyton on omnivore density. Both models suggested a positive relationship between small herbivores and small omnivores, indicating that predation was unable to limit herbivore numbers. Total effects of time following flooding were negative for all three consumer groups even when both preferred models suggested positive direct effects for some groups. Total effects of nutrient levels (phosphorus) were positive for consumers and generally larger than those of hydrological disturbance and were mediated by changes in periphyton content. 4. Our findings provide quantitative support for indirect effects of nutrient enrichment on consumers, and the importance of both algal community structure and periphyton biomass to Everglades food webs. Evidence for top‐down control of infauna by omnivores was noted, though without substantially greater support than a competing bottom‐up‐only model.     

Fish predation regimes modify benthic diatom community structures: Experimental evidence from an in situ mesocosm study

Diatoms are important primary producers in shallow water environments. Few studies have assessed the importance of biological interactions in structuring these communities. In the present study, benthic diatom community structure in relation to manipulated food webs was assessed using in situ mesocosms, whereby predator‐free environments and environments comprising two different fish species were assessed. Zooplankton abundance, settled algal biomass and the diatom community were monitored over a 12‐day period across each of the three trophic scenarios. Differences among treatments over time were observed in zooplankton abundances, particularly copepods. Similarly, the benthic diatom community structure changed significantly over time across the three trophic treatments. However, no differences in total algal biomass were found among treatments. This was likely the result of non‐diatom phytoplankton contributions. We propose that the benthic diatom community structure within the mesocosms was influenced by trophic cascades and potentially through direct consumption by the fish. The study highlights that not only are organisms at the base of the food web affected by predators at the top of the food web, but that predator identity is potentially an important consideration for predator–prey interaction outcomes with consequences for multiple trophic levels.     

Influence of food‐web structure on the biodegradability of lake sediment

1. Sediment plays a key role in internal nutrient cycling and eutrophication in lakes. However, studies focusing on the efficiency of the biomanipulation techniques for improving the control of primary producers have rarely examined the effects of changes in food‐web structure on the sediment biochemical composition and biodegradability. 2. In a 1‐year experiment conducted in large replicated mesocosms, we tested how the absence or presence of a zooplanktivorous fish (roach, Rutilus rutilus) affected the elemental composition and the potential biodegradability of recently deposited sediment in a eutrophic system. The potential biodegradability of these sediments was assessed in laboratory microcosms by measuring the production of CO₂ during 44‐day incubations. 3. The potential biodegradability of recently deposited sediment from the fish treatment was 60% higher than that from the fishless treatment. This higher biodegradability was corroborated by a higher annual loss of sediment in fish enclosures (36%) than in fishless ones (16%). Annual losses of carbon, nitrogen and organic phosphorous were higher for sediment from fish enclosures. 4. Carbon and nitrogen contents of sediment were higher for the fish treatment. In contrast, the sediment C/N ratio, one of the proxies used to estimate sediment biodegradability, did not differ between treatments. No relationship was observed between elemental composition of sediment and its potential biodegradability. This latter appeared to be more probably dependent on the biochemical composition of the sediment and especially on the content of labile compounds such as proteins, sugars and polyunsaturated fatty acids. The use of sterols as biomarkers revealed an important degradation by microorganisms of 1‐year‐old sediment from both fish and fishless treatments. 5. Our results revealed that fish biomanipulations might favour clear water states not only through a stronger top–down control on phytoplankton but also through a lower biodegradability of sediment reducing internal nutrient cycling.     

Warming shifts top-down and bottom-up control of pond food web structure and function

The effects of global and local environmental changes are transmitted through networks of interacting organisms to shape the structure of communities and the dynamics of ecosystems. We tested the impact of elevated temperature on the top-down and bottom-up forces structuring experimental freshwater pond food webs in western Canada over 16 months. Experimental warming was crossed with treatments manipulating the presence of planktivorous fish and eutrophication through enhanced nutrient supply. We found that higher temperatures produced top-heavy food webs with lower biomass of benthic and pelagic producers, equivalent biomass of zooplankton, zoobenthos and pelagic bacteria, and more pelagic viruses. Eutrophication increased the biomass of all organisms studied, while fish had cascading positive effects on periphyton, phytoplankton and bacteria, and reduced biomass of invertebrates. Surprisingly, virus biomass was reduced in the presence of fish, suggesting the possibility for complex mechanisms of top-down control of the lytic cycle. Warming reduced the effects of eutrophication on periphyton, and magnified the already strong effects of fish on phytoplankton and bacteria. Warming, fish and nutrients all increased whole-system rates of net production despite their distinct impacts on the distribution of biomass between producers and consumers, plankton and benthos, and microbes and macrobes. Our results indicate that warming exerts a host of indirect effects on aquatic food webs mediated through shifts in the magnitudes of top-down and bottom-up forcing.     

Convergence of trophic interaction strengths in grassland food webs through metabolic scaling of herbivore biomass

1. Food web theory hypothesizes that trophic interaction strengths of consumers should vary with consumer metabolic body mass (mass(0·75) ) rather than simply with consumer body mass (mass(1·0) ) owing to constraints on consumption imposed by metabolic demand for and metabolic capacity to process nutrients and energy. Accordingly, species with similar metabolic body masses should have similar trophic interaction strengths. 2. We experimentally tested this hypothesis by assembling food webs comprised of species of arthropod predators, small sap-feeding and large leaf-chewing insect herbivores and herbaceous plants in a New England, USA meadow grassland. The experiment comprised of a density-matching treatment where herbivore species were stocked into field mesocosms at equal densities to quantify baseline species identity and metabolic body mass effects. The experiment also comprised of a metabolic biomass-matching treatment where smaller sap-feeding herbivore (SH) species were stocked into mesocosms such that the product of their density and metabolic body mass (metabolic biomass) was equal to the large herbivore (LH) species. We compared the magnitude of the direct effects of herbivore species on plants in the different treatments. We also compared the magnitude of indirect effects between predators and plants mediated by herbivores in the different treatments. 3. Consistent with the hypothesis, we found that increasing metabolic biomass translated into a 9-14-fold increase in magnitude of herbivore direct effects and up to a fivefold increase in indirect effects on plants. Moreover, metabolic biomass matching caused interaction strengths among herbivore species to converge. This result came about through increases in the herbivore mean effects as well as decreases in variation in effects among treatment replicates as herbivore metabolic biomass increased. 4. We found, however, that herbivore feeding mode rather than herbivore metabolic biomass explained differences in the sign of indirect effects in the different food webs. 5. We conclude that increasing herbivore metabolic biomass not only strengthened the direct and indirect effects on plants but also made those effects more consistent across space. Nevertheless, metabolic biomass alone could not completely explain variation in the nature of indirect effects in the food web, suggesting that additional consideration of consumer traits like feeding mode will provide a more nuanced understanding of trophic interaction strengths in food webs.     

Snail–periphyton interactions in a prairie lacustrine wetland

This study examined the effects of nutrients and macrophytes on snail grazers and periphyton in a prairie wetland food web. Snails (Gyraulus circumstriatus) and periphyton in large enclosures in a lacustrine wetland, Delta Marsh, MB, Canada were subjected to two experimental treatments, nutrient addition (nitrogen, phosphorus) and macrophyte exclusion (using a porous geotextile carpet) during July and August. Snail biomass and periphyton biomass (on both artificial substrata and submerged macrophytes) increased over time in all treatments, representing seasonal growth. Snail biomass was three times higher on macrophytes than on artificial substrata. In response to nutrient addition, snail biomass was significantly elevated over time on macrophytes but not on artificial substrata. Conversely, periphyton biomass was higher on artificial substrata but not on macrophytes in response to nutrient addition. Snail biomass and periphyton biomass on artificial substrata showed no response to macrophyte exclusion. Snail biomass on all substrata was inversely correlated with turbidity, whereas periphyton biomass showed no relationship with turbidity. Timing of nutrient additions to wetlands may influence whether the response occurs primarily in phytoplankton or in periphyton and macrophytes.     

Indirect food web interactions increase growth of an algivorous stream fish

1. We tested the hypothesis that indirect food web interactions between some common, invertivorous fishes and their prey would positively affect growth of an algivorous fish species. Specifically, we predicted that orangethroat darter (Etheostoma spectabile) would increase periphyton biomass via a top‐down pathway, indirectly enhancing growth of the algivorous central stoneroller minnow (Campostoma anomalum). Moreover, we predicted that sand shiner (Notropis stramineus) would increase periphyton biomass via a bottom‐up pathway and indirectly enhance growth of the stoneroller minnow. 2. In an 83‐day experiment in large, outdoor, stream mesocosms, we stocked two fish species per mesocosm (stoneroller and either darter or shiner), estimated the effects of the invertivorous and grazing fishes on periphyton biomass and estimated growth of the algivorous fish. 3. The darter consumed grazing invertebrates, indirectly increasing periphyton biomass. The shiner consumed terrestrial insects as predicted, but it did not affect periphyton biomass. 4. In support of our hypothesis, the darter indirectly enhanced stoneroller growth. As predicted, stonerollers consumed the increased periphyton in streams with darters, resulting in greater growth, condition and gut fullness compared to streams without darters. No indirect interaction was observed between stonerollers and shiners. 5. Our study suggests that some invertivorous fish species can positively affect growth of algivorous fishes through indirect food web interactions. Thus, in stream communities, it is possible that the loss of a single, invertivorous fish taxon could have negative consequences on algivorous fish populations via the removal of positive indirect food web interactions.     

Elevated light and nutrients alter the nutritional quality of stream periphyton

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High nutrient availability leads to weaker top‐down control of stream periphyton: Compensatory feeding in Ancylus fluviatilis

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Light intensity regulates growth and reproduction of a snail grazer (Gyraulus chinensis) through changes in the quality and biomass of stream periphyton

1. The light : nutrient hypothesis (LNH) proposes that herbivore growth rates are maximised at intermediate light‐to‐nutrient ratios. A reduction to light intensity (i.e. decreased light‐to‐nutrient ratio) should lead to reduced food availability for herbivores while excessive light intensity in oligotrophic environments (i.e. increased light‐to‐nutrient ratios) should increase the C : N and C : P ratios of producers. However, this hypothesis has not yet been supported by studies on stream ecosystems. 2. We tested the LNH by experimental application of controlled natural gradients in light intensity to oligotrophic laboratory channels that included periphyton and the freshwater snail Gyraulus chinensis. 3. The results in this oligotrophic environment indicate that light regulated the flow of matter between trophic levels and grazer reproduction by controlling C : P ratios of the producers.     

The influence of water level on macrophyte growth and trophic interactions in eutrophic Mediterranean shallow lakes: a mesocosm experiment with and without fish

1. Water‐level fluctuations are typical of lakes located in the semi‐arid Mediterranean region, which is characterised by warm rainy winters and hot dry summers. Ongoing climate change may exacerbate fluctuations and lead to more severe episodes of drought, so information on the effects of water level on the functioning of lake ecosystems in such regions is crucial. 2. In eutrophic Lake Eymir, Turkey, we conducted a 4‐month (summer) field experiment using cylindrical 0.8‐m‐ (low‐water‐level) and 1.6‐m‐deep (high‐water‐level) mesocosms (kept open to the sediment and atmosphere). Fish (tench, Tinca tinca, and bleak, Alburnus escherichii) were added to half of the mesocosms, while the rest were kept fishless. Ten shoots of Potamogeton pectinatus were transplanted to each mesocosm. 3. Sampling for physicochemical variables, chlorophyll a (chl‐a), zooplankton and per cent plant volume inhabited (PVI%) by macrophytes was conducted weekly during the first 5 weeks, and subsequently biweekly. Macrophytes were harvested on the last sampling date. During the course of the experiment, the water level decreased by 0.41 ± 0.06 m. 4. Throughout the experiment, fish affected zooplankton abundance (?), nutrient concentrations (+), chl‐a (+) and water clarity (?) most strongly in the low‐water‐level mesocosms and the zooplankton community shifted towards dominance of small‐sized forms. The fishless mesocosms had a higher zooplankton/phytoplankton ratio, suggesting higher grazing. 5. Greatest macrophyte growth was observed in the low‐water‐level fishless mesocosms. However, despite high nutrient concentrations and low water clarity, macrophytes were also abundant in the fish mesocosms and particularly increased following a water‐level decrease from midsummer onwards. Macrophyte growth was poor in the high‐water‐level mesocosms, even in the fishless ones with high water clarity. This was ascribed to extensive periphyton development reducing light availability for the macrophytes. 6. Our results indicate that a reduction in water level during summer may help maintain the growth of macrophytes in Mediterranean eutrophic shallow lakes, despite a strong negative effect of fish predation on water clarity. It is therefore probable that an expected negative effect of global climate change on water clarity because of eutrophication and enhanced top‐down control of fish may be, at least partly, counteracted by reduced water level, provided that physical disturbance is not severe.     

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

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Nutrient loading and grazing by the minnow Phoxinus erythrogaster shift periphyton abundance and stoichiometry in mesocosms

1. Anthropogenic activities in prairie streams are increasing nutrient inputs and altering stream communities. Understanding the role of large consumers such as fish in regulating periphyton structure and nutritional content is necessary to predict how changing diversity will interact with nutrient enrichment to regulate stream nutrient processing and retention. 2. We characterised the importance of grazing fish on stream nutrient storage and cycling following a simulated flood under different nutrient regimes by crossing six nutrient concentrations with six densities of a grazing minnow (southern redbelly dace, Phoxinus erythrogaster) in large outdoor mesocosms. We measured the biomass and stoichiometry of overstory and understory periphyton layers, the stoichiometry of fish tissue and excretion, and compared fish diet composition with available algal assemblages in pools and riffles to evaluate whether fish were selectively foraging within or among habitats. 3. Model selection indicated nutrient loading and fish density were important to algal composition and periphyton carbon (C): nitrogen (N). Nutrient loading increased algal biomass, favoured diatom growth over green algae and decreased periphyton C : N. Increasing grazer density did not affect biomass and reduced the C : N of overstory, but not understory periphyton. Algal composition of dace diet was correlated with available algae, but there were proportionately more diatoms present in dace guts. We found no correlation between fish egestion/excretion nutrient ratios and nutrient loading or fish density despite varying N content of periphyton. 4. Large grazers and nutrient availability can have a spatially distinct influence at a microhabitat scale on the nutrient status of primary producers in streams.     

Feeding by omnivores increases food available to consumers

Consumers are usually thought of as negatively affecting producers, but they can affect them positively by releasing nutrients (nutrient regeneration). The net effects of consumers on producers should depend on the balance between the effects of consumption and nutrient regeneration. In aquatic habitats, nutrient regeneration by consumers may increase microbial activity on leaf detritus as well as algal production, which in turn may stimulate further nutrient release and benefit herbivores or detritivores by increasing food quantity or quality. Omnivores can regenerate nutrients from animals, algae and detritus, creating diverse nutrient pathways. Many tadpoles are omnivores, and their nutrient regeneration may be important in aquatic food webs. To reveal the nutrient pathways created by tadpoles and examine whether omnivorous tadpoles can have positive effects on producers and consumers, we experimentally examined the effects of nutrient regeneration by three densities of tadpoles on primary producers, leaf litter, and other consumers in tank mesocosms. Tadpole exclosures were placed inside each mesocosm, allowing us to separate direct consumption effects from indirect nutrient regeneration effects. Nutrient regeneration caused by the herbivorous and carnivorous feeding activities of tadpoles positively affected rates of production of benthic algae, phytoplankton, and herbivorous benthic chironomid larvae, and rates of mineralization of leaf litter. The increased production of benthic algae and chironomid larvae was consumed by the tadpoles themselves, leaving no net change in the standing biomass of these resources. Our experiment thus demonstrated that omnivores created complicated nutrient pathways and accelerated rates of primary production and growth rates of other consumers, leading to increased rates of food availability to the omnivores themselves. Interactions of this nature may be common in many systems and could strongly moderate the effects of consumers on their resources and each other.     

Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.