Climate‐mediated changes in marine ecosystem regulation during El Niño

The degree to which ecosystems are regulated through bottom‐up, top‐down, or direct physical processes represents a long‐standing issue in ecology, with important consequences for resource management and conservation. In marine ecosystems, the role of bottom‐up and top‐down forcing has been shown to vary over spatio‐temporal scales, often linked to highly variable and heterogeneously distributed environmental conditions. Ecosystem dynamics in the Northeast Pacific have been suggested to be predominately bottom‐up regulated. However, it remains unknown to what extent top‐down regulation occurs, or whether the relative importance of bottom‐up and top‐down forcing may shift in response to climate change. In this study, we investigate the effects and relative importance of bottom‐up, top‐down, and physical forcing during changing climate conditions on ecosystem regulation in the Southern California Current System (SCCS) using a generalized food web model. This statistical approach is based on nonlinear threshold models and a long‐term data set (~60 years) covering multiple trophic levels from phytoplankton to predatory fish. We found bottom‐up control to be the primary mode of ecosystem regulation. However, our results also demonstrate an alternative mode of regulation represented by interacting bottom‐up and top‐down forcing, analogous to wasp‐waist dynamics, but occurring across multiple trophic levels and only during periods of reduced bottom‐up forcing (i.e., weak upwelling, low nutrient concentrations, and primary production). The shifts in ecosystem regulation are caused by changes in ocean‐atmosphere forcing and triggered by highly variable climate conditions associated with El Niño. Furthermore, we show that biota respond differently to major El Niño events during positive or negative phases of the Pacific Decadal Oscillation (PDO), as well as highlight potential concerns for marine and fisheries management by demonstrating increased sensitivity of pelagic fish to exploitation during El Niño.     

Bottom‐up vs. top‐down effects on terrestrial insect herbivores: a meta‐analysis

Primary consumers are under strong selection from resource (‘bottom‐up’) and consumer (‘top‐down’) controls, but the relative importance of these selective forces is unknown. We performed a meta‐analysis to compare the strength of top‐down and bottom‐up forces on consumer fitness, considering multiple predictors that can modulate these effects: diet breadth, feeding guild, habitat/environment, type of bottom‐up effects, type of top‐down effects and how consumer fitness effects are measured. We focused our analyses on the most diverse group of primary consumers, herbivorous insects, and found that in general top‐down forces were stronger than bottom‐up forces. Notably, chewing, sucking and gall‐making herbivores were more affected by top‐down than bottom‐up forces, top‐down forces were stronger than bottom‐up in both natural and controlled (cultivated) environments, and parasitoids and predators had equally strong top‐down effects on insect herbivores. Future studies should broaden the scope of focal consumers, particularly in understudied terrestrial systems, guilds, taxonomic groups and top‐down controls (e.g. pathogens), and test for more complex indirect community interactions. Our results demonstrate the surprising strength of forces exerted by natural enemies on herbivorous insects, and thus the necessity of using a tri‐trophic approach when studying insect‐plant interactions.     

Structural equation models suggest that bottom‐up processes override top‐down processes in boreal pikeperch (Sander lucioperca) lakes

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Density‐ and trait‐mediated top–down effects modify bottom–up control of a highly endemic tropical aquatic food web

Benthic invertebrates mediate bottom–up and top–down influences in aquatic food webs, and changes in the abundance or traits of invertebrates can alter the strength of top–down effects. Studies assessing the role of invertebrate abundance and behavior as controls on food web structure are rare at the whole ecosystem scale. Here we use a comparative approach to investigate bottom–up and top–down influences on whole anchialine pond ecosystems in coastal Hawai‘i. In these ponds, a single species of endemic atyid shrimp (Halocaridina rubra) is believed to structure epilithon communities. Many Hawaiian anchialine ponds and their endemic fauna, however, have been greatly altered by bottom–up (increased nutrient enrichment) and top–down (introduced fish predators) disturbances from human development. We present the results of a survey of dissolved nutrient concentrations, epilithon biomass and composition, and H. rubra abundance and behavior in anchialine ponds with and without invasive predatory fish along a nutrient concentration gradient on the North Kona coast of Hawai‘i. We use linear models to assess 1) the effects of nutrient loading and fish introductions on pond food web structure and 2) the role of shrimp density and behavior in effecting that change. We find evidence for bottom–up food web control, in that nutrients were associated with increased epilithon biomass, autotrophy and nutrient content as well as increased abundance and size of H. rubra. We also find evidence for top–down control, as ponds with invasive predatory fish had higher epilithon biomass, productivity, and nutrient content. Top–down effects were transmitted by both altered H. rubra abundance, which changed the biomass of epilithon, and H. rubra behavior, which changed the composition of the epilithon. Our study extends experimental findings on bottom–up and top–down control to the whole ecosystem scale and finds evidence for qualitatively different effects of trait‐ and density‐mediated change in top–down influences.     

Seasonal,inter‐annual and long term variation in top–down versus bottom–up regulation of primary production

Environmental change strongly affects primary productivity of ecosystems via modifying bottom–up and top–down regulation of primary producers. Here we present a novel approach to quantify the relative importance of regulating factors in natural systems over various time scales: we calculated daily effect sizes of major factors affecting phytoplankton growth during the spring bloom period during almost three decades of lake oligotrophication using numerical experiments with a data based simulation model. We show that with oligotrophication the regulation of spring phytoplankton shifts from primarily top–down to bottom–up, and that the changes in regulation are non‐linearly related to the nutrient (phosphorus) concentrations. Our findings indicate that long‐term changes in top–down regulation cannot be understood without considering multiple herbivore taxa, here, microzooplankton (ciliates) and mesozooplankton (daphnids). We further demonstrate that bottom–up and top–down regulation are not independent from each other and that their interaction is time‐scale dependent.     

Up or down in space? Uniting the bottom‐up versus top‐down paradigm and spatial ecology

Why is the World green – what keeps herbivores, and herbivorous insects in particular, from consuming all of their food? When this question was first posed, the relative importance of top‐down and bottom‐up effects was hotly disputed. While modern ecologists may agree that impacts from several different directions will affect local insect densities, the bottom‐up vs top‐down jargon seems to be stuck in a unidimensional world. Here, we argue that the strength of almost every bottom‐up and top‐down force is likely to vary in space, and that in itself, spatial structure invokes new processes which defy classification in the traditional bottom‐up top‐down scheme. To understand the relative importance of different forces keeping herbivore numbers in check, we feel that we need a fresh synthesis between the novel paradigm of spatial ecology and the classical paradigms of top‐down and bottom‐up studies. This synthesis requires a consideration of forces beyond the standard framework of top‐down vs bottom‐up effects, and should be based on comparing the relative strength of such forces at several sites in a spatially explicit framework. Overall, we should switch our focus from whether the relative strength of top‐down and bottom‐up factors vary in space to why there is variation, how much variation there is, and at what spatial scale(s) it occurs.     

Hybrid Framework for Managing Uncertainty in Life Cycle Inventories

Life cycle assessment (LCA) is increasingly being used to inform decisions related to environmental technologies and polices, such as carbon footprinting and labeling, national emission inventories, and appliance standards. However, LCA studies of the same product or service often yield very different results, affecting the perception of LCA as a reliable decision tool. This does not imply that LCA is intrinsically unreliable; we argue instead that future development of LCA requires that much more attention be paid to assessing and managing uncertainties. In this article we review past efforts to manage uncertainty and propose a hybrid approach combining process and economic input–output (I‐O) approaches to uncertainty analysis of life cycle inventories (LCI). Different categories of uncertainty are sometimes not tractable to analysis within a given model framework but can be estimated from another perspective. For instance, cutoff or truncation error induced by some processes not being included in a bottom‐up process model can be estimated via a top‐down approach such as the economic I‐O model. A categorization of uncertainty types is presented (data, cutoff, aggregation, temporal, geographic) with a quantitative discussion of methods for evaluation, particularly for assessing temporal uncertainty. A long‐term vision for LCI is proposed in which hybrid methods are employed to quantitatively estimate different uncertainty types, which are then reduced through an iterative refinement of the hybrid LCI method.     

A unified treatment of top-down and bottom-up control of reproduction in populations

Generalizations describing how top‐down and bottom‐up processes jointly influence the production of offspring (recruitment) and the number of reproducing adults are lacking. This is a deficiency because (1) it is widely recognized that both top‐down and bottom‐up processes are common in ecosystems; and (2) the relationship between the number of individuals recruiting and number of reproductively active individuals present in that population is of fundamental importance in all branches of ecology. Here we derive a model to consider the joint effects of top‐down and bottom‐up forcing in any ecosystem. In general, during the lifetime of a cohort, bottom‐up effects are likely to limit recruitment over longer periods of time than top‐down effects. Top‐down effects are likely to be most important early in the life history when potential recruits are small in size, and such effects will be more recognizable in small cohorts comprised of slowly growing individuals.     

Energy simulation and LCA for macro-scale analysis of eco-innovations in the housing stock

Purpose

Energy consumption of buildings is one of the major drivers of environmental impacts. Life cycle assessment (LCA) may support the assessment of burdens and benefits associated to eco-innovations aiming at reducing these environmental impacts. Energy efficiency policies however typically focus on the meso- or macro-scale, while interventions are typically taken at the micro-scale. This paper presents an approach that bridges this gap by using the results of energy simulations and LCA studies at the building level to estimate the effect of micro-scale eco-innovations on the macro-scale, i.e. the housing stock in Europe.

Methods

LCA and dynamic energy simulations are integrated to accurately assess the life cycle environmental burdens and benefits of eco-innovation measures at the building level. This allows quantitatively assessing the effectiveness of these measures to lower the energy use and environmental impact of buildings. The analysis at this micro-scale focuses on 24 representative residential buildings within the EU. For the upscaling to the EU housing stock, a hybrid approach is used. The results of the micro-scale analysis are upscaled to the EU housing stock scale by adopting the eco-innovation measures to (part of) the EU building stock (bottom–up approach) and extrapolating the relative impact reduction obtained for the reference buildings to the baseline stock model. The reference buildings in the baseline stock model have been developed by European Commission-Joint Research Centre based on a statistical analysis (top–down approach) of the European housing stock. The method is used to evaluate five scenarios covering various aspects: building components (building envelope insulation), technical installations (renewable energy), user behaviour (night setback of the setpoint temperature), and a combined scenario.

Results and discussion

Results show that the proposed combination of bottom–up and top–down approaches allow accurately assessing the impact of eco-innovation measures at the macro-scale. The results indicate that a combination of policy measures is necessary to lower the environmental impacts of the building stock to a significative extent.

Conclusions

Interventions addressing energy efficiency at building level may lead to the need of a trade-off between resource efficiency and environmental impacts. LCA integrated with dynamic energy simulation may help unveiling the potential improvements and burdens associated to eco-innovations.

     

The paradox of re‐oligotrophication: the role of bottom–up versus top–down controls on the phytoplankton community

Increases in phytoplankton biomass have been widely observed over the past decades, even in lakes experiencing nutrient reduction. However, the mechanisms giving rise to this trend remain unclear. Here, we unveil the potential mechanisms through quantifying the relative contribution of bottom–up versus top–down control in determining biomasses of phytoplankton assemblages in Lake Geneva. Specifically, we apply nonlinear time series analysis, convergent cross mapping (CCM), to decipher the degree of bottom–up versus top–down control among phytoplankton assemblages via quantifying 1) causal links between environmental factors and various phytoplankton assemblages and 2) the relative importance of bottom–up, top–down, and environmental effects. We show that the recent increase in total phytoplankton biomass, albeit with phosphorus reduction, was mainly caused by a particular phytoplankton assemblage which was better adapted to the re‐oligotrophicated environment characterized by relatively low phosphorus concentrations and warm water temperature, and poorly controlled by zooplankton grazing. Our findings suggest that zooplankton act as a critical driver of phytoplankton biomass and strongly impact the dynamics of recovery from eutrophication. Therefore, our phytoplankton assemblage approach in combination with causal identification of top–down versus bottom–up controls provides insights into the reason why phytoplankton biomass may increase in lakes undergoing phosphorus reduction.     

A Methodical Approach for Systematic Life Cycle Assessment of Wood‐Based Furniture

Existing life cycle assessment (LCA) studies for furniture focus on single pieces of furniture and use a bottom‐up approach based on their bill of materials (BOM) to build up the data inventories. This approach does not ensure completeness regarding material and energy fluxes and representativeness regarding the product portfolio. Integrating material and energy fluxes collected at company level into product LCA (top‐down approach) over‐rides this drawback. This article presents a method for systematic LCA of industrially produced furniture that merges the top‐down approach and bottom‐up approach. The developed method assigns data collected at the company level to the different products while, at the same time, considering that wood‐based furniture is a complex product. Hence, several classifications to reduce the complexity to a manageable level have been developed. Simultaneously, a systematic calculation routine was established. The practical implementation of the developed method for systematic LCA is carried out in a case study within the German furniture industry. The system boundary was set in accord with the EN 15804 specification cradle‐to‐gate‐with‐options. The analysis therefore includes the manufacturing phase supplemented by an end‐of‐life scenario. The case study shows that the manufacturing of semifinished products (especially wood‐based panels and metal components) as well as the electric energy demand in furniture manufacturing account for a notable share of the environmental impacts. A sensitivity analysis indicates that up to roughly 10% of the greenhouse gas emissions are not recorded when conducting an LCA based on a BOM instead of applying the developed approach.     

Effects of plant quality and ant defence on herbivory rates in a neotropical ant‐plant

1. Understanding the degree to which populations and communities are limited by both bottom‐up and top‐down effects is still a major challenge for ecologists, and manipulation of plant quality, for example, can alter herbivory rates in plants. In addition, biotic defence by ants can directly influence the populations of herbivores, as demonstrated by increased rates of herbivory or increased herbivore density after ant exclusion. The aim of this study was to evaluate bottom‐up and top‐down effects on herbivory rates in a mutualistic ant‐plant. 2. In this study, the role of Azteca alfari ants as biotic defence in individuals of Cecropia pachystachya was investigated experimentally with a simultaneous manipulation of both bottom‐up (fertilisation) and top‐down (ant exclusion) factors. Four treatments were used in a fully factorial design, with 15 replicates for each treatment: (i) control plants, without manipulation; (ii) fertilised plants, ants not manipulated; (iii) unfertilised plants and excluded ants and (iv) fertilised plants and ants excluded. 3. Fertilisation increased the availability of foliar nitrogen in C. pachystachya, and herbivory rates by chewing insects were significantly higher in fertilised plants with ants excluded. 4. Herbivory, however, was more influenced by bottom‐up effects – such as the quality of the host plant – than by top‐down effects caused by ants as biotic defences, reinforcing the crucial role of leaf nutritional quality for herbivory levels experienced by plants. Conditionality in ant defence under increased nutritional quality of leaves through fertilisation might explain increased levels of herbivory in plants with higher leaf nitrogen.     

How to measure top–down vs bottom–up effects: a new population metric and its calibration on Daphnia

Research on the role of top–down (predation) and bottom–up (food) effects in food webs has led to the understanding that the variability of these effects in space and time is a fundamental feature of natural systems. Consequently, our measurement tools must allow us to evaluate the effects from a dynamical perspective. A population‐dynamics approach may be appropriate to the task. More specifically, because food and predators both affect birth rate, birth rate dynamics may be a key to understanding their impact on the population of interest. Based on the Edmondson–Paloheimo model for birth rate, we propose a new population metric to assess the relative strength of top–down vs bottom–up effects. The metric is the ratio of contributions of changes in proportion of adults and fecundity to change in birth rate. Proportion of adults reflects a top–down effect (predators are assumed to be size‐selective), fecundity reflects a bottom–up effect, and birth rate appears as a common currency with which to compare the former and the latter. Using microcosm experiments and computer simulations on the cladoceran Daphnia, we calibrate the metric and show that, in both types of tests, the ratio of contributions is typically 0.5–0.7 under a strong bottom–up effect and 2.0–2.2 under a strong top–down effect. This provides experimental evidence that the ratio of contributions may allow one to distinguish a strong top–down effect from a strong bottom–up effect.     

Spatial variability of the plankton trophic interaction in the North Sea: a new feature after the early 1970s

Traditionally, marine ecosystem structure was thought to be bottom‐up controlled. In recent years, a number of studies have highlighted the importance of top‐down regulation. Evidence is accumulating that the type of trophic forcing varies temporally and spatially, and an integrated view – considering the interplay of both types of control – is emerging. Correlations between time series spanning several decades of the abundances of adjacent trophic levels are conventionally used to assess the type of control: bottom‐up if positive or top‐down if this is negative. This approach implies averaging periods which might show time‐varying dynamics and therefore can hide part of this temporal variability. Using spatially referenced plankton information extracted from the Continuous Plankton Recorder, this study addresses the potential dynamic character of the trophic structure at the planktonic level in the North Sea by assessing its variation over both temporal and spatial scales. Our results show that until the early‐1970s a bottom‐up control characterized the base of the food web across the whole North Sea, with diatoms having a positive and homogeneous effect on zooplankton filter‐feeders. Afterwards, different regional trophic dynamics were observed, in particular a negative relationship between total phytoplankton and zooplankton was detected off the west coast of Norway and the Skagerrak as opposed to a positive one in the southern reaches. Our results suggest that after the early 1970s diatoms remained the main food source for zooplankton filter‐feeders east of Orkney–Shetland and off Scotland, while in the east, from the Norwegian Trench to the German Bight, filter‐feeders were mainly sustained by dinoflagellates.     

Top‐down,bottom‐up,or side to side? Within‐trophic‐level interactions modify trophic dynamics of a salt marsh herbivore

Many factors can influence the top‐down and bottom‐up dynamics of phytophagous insects. Although interactions between herbivore species have been frequently shown to be ecologically important, the effects of such horizontal trophic interactions on the relative roles of top‐down and bottom‐up forces have gone largely unstudied. In this paper we report on the results of a factorial field experiment in which we examined the effects of within‐trophic‐level interactions on the top‐down and bottom‐up dynamics of a salt marsh planthopper.
We manipulated the bottom‐up effects of plant quality by increasing soil salinity, and manipulated top‐down effects by decreasing the intensity of parasitoid attack with yellow sticky traps that removed hymenopteran parasitoids. We applied these treatments to plots in two patches of the host plant, one with low densities of lepidopteran stem borer larvae, and one with high densities of stem borers. We maintained the treatments and monitored planthopper density for ten months, from March through December 1999. Increased salinity significantly increased planthopper density within one month of the first application of salt. The rapid response of the planthopper to salt treatments suggested a chemical mechanism, perhaps mobilization of bound nitrogen. Yellow sticky traps, although significantly reducing parasitism of planthopper eggs, had little impact on hopper density. The density of lepidopteran stem borers, however, had an even greater impact on planthopper density than did salt treatments, with high stem borer plots supporting much lower densities of hoppers. Stem borer density also reduced the response of the planthopper to other treatments, especially salt supplementation. The results of this study show that the impact of within‐trophic‐level interactions can significantly change herbivore trophic dynamics and can be even more important than either top‐down or bottom‐up effects in determining herbivore density.     

Bottom‐up effects of glucosinolate variation on aphid colony dynamics in wild cabbage populations

1. There is an ongoing debate about the relative importance of top‐down and bottom‐up regulation of herbivore dynamics in the wild. Secondary metabolites, produced by plants, have negative effects on survival and growth of some herbivore species, causing bottom‐up regulation of population dynamics. Herbivore natural enemies may use plant secondary metabolites as cues to find their prey, but their survival and reproduction can also be influenced by the upward cascade of secondary metabolites through the food web. Thus plant chemistry might also affect herbivore populations by mediating top‐down regulation. 2. We investigated the influence of heritable variation in aliphatic glucosinolates, a class of secondary metabolites produced by Brassica plants, on the relative importance of top‐down and bottom‐up regulation of Brevicoryne brassicae (mealy cabbage aphid) colonies in natural Brassica oleracea (wild cabbage) populations. We manipulated natural enemy pressure on plants differing in their glucosinolate profiles, and monitored aphid colony growth and disperser production. 3. Aphid colony sizes were significantly smaller on plants producing sinigrin, compared with plants producing alternative aliphatic glucosinolates. Aphid natural enemy numbers correlated with aphid colony size, but there was no additional effect of the plants' chemical phenotype on natural enemy abundance. Furthermore, experimental reduction of natural enemy pressure had no effect on aphid colony size or production of winged dispersers. 4. Our results provide evidence for glucosinolate‐mediated, bottom‐up regulation of mealy cabbage aphid colonies in natural populations, but we found no indication of top‐down regulation. We emphasise that more studies of these processes should focus on tritrophic interactions in the wild.     

Bottom–up and top–down effects on plant communities: nutrients limit productivity,but insects determine diversity and composition

Top–down effects of herbivores and bottom–up effects of nutrients shape productivity and diversity across ecosystems, yet their single and combined effects on spatial and temporal beta diversity is unknown. We established a field experiment in which the abundance of insect herbivores (top–down) and soil nitrogen (bottom–up) were manipulated over six years in an existing old‐field community. We tracked plant α and β diversity – within plot richness and among plot biodiversity‐ and aboveground net primary productivity (ANPP) over the course of the experiment. We found that bottom–up factors affected ANPP while top–down factors influenced plant community structure. Across years, while N reduction lowered ANPP by 10%, N reduction did not alter ANPP relative to control plots. Further, N reduction lowered ANPP by 20% relative to N addition plots. On the other hand, the reduction of insect herbivores did not alter plant richness (α diversity) yet consistently promoted Shannon's evenness, relative to plots where insect herbivores were present. Further, insect herbivores promoted spatial‐temporal β diversity. Overall, we found that the relative importance of top–down and bottom–up controls of plant ANPP, plant α diversity, and composition (β diversity) can vary significantly in magnitude and direction. In addition, their effects varied through time, with bottom–up effects influencing ANPP quickly while the effects of top–down factors emerging only late in the experiment to influence plant community composition via shifts in plant dominance.     

Bottom‐up processes drive reproductive success in an apex predator

One of the central goals of the field of population ecology is to identify the drivers of population dynamics, particularly in the context of predator–prey relationships. Understanding the relative role of top‐down versus bottom‐up drivers is of particular interest in understanding ecosystem dynamics. Our goal was to explore predator–prey relationships in a boreal ecosystem in interior Alaska through the use of multispecies long‐term monitoring data. We used 29 years of field data and a dynamic multistate site occupancy modeling approach to explore the trophic relationships between an apex predator, the golden eagle, and cyclic populations of the two primary prey species available to eagles early in the breeding season, snowshoe hare and willow ptarmigan. We found that golden eagle reproductive success was reliant on prey numbers, but also responded prior to changes in the phase of the snowshoe hare population cycle and failed to respond to variation in hare cycle amplitude. There was no lagged response to ptarmigan populations, and ptarmigan populations recovered quickly from the low phase. Together, these results suggested that eagle reproduction is largely driven by bottom‐up processes, with little evidence of top‐down control of either ptarmigan or hare populations. Although the relationship between golden eagle reproductive success and prey abundance had been previously established, here we established prey populations are likely driving eagle dynamics through bottom‐up processes. The key to this insight was our focus on golden eagle reproductive parameters rather than overall abundance. Although our inference is limited to the golden eagle–hare–ptarmigan relationships we studied, our results suggest caution in interpreting predator–prey abundance patterns among other species as strong evidence for top‐down control.     

Long Term Effect of Cyprinid Fishes on Phytoplankton and Zooplankton Communities in a Shallow Water Protection Reservoir

The effects of fish kill and different fish stocks on the phytoplankton and zooplankton dynamics were studied in a shallow hypertrophic reservoir system. When fish stock was below 100 kg ha⁻¹, nutrient availability was not the main limiting factor for growth of phytoplankton. Consequently top‐down forces controlled phytoplankton. In the years with high fish stock (>100 kg ha⁻¹) the bottom‐up forces dominated as nutrient availability was the main limiting factor for growth of phytoplankton. We can conclude that significant water quality improvement can be achieved in the reservoir system by decreasing fish stock below 100 kg ha⁻¹. Although clear‐water phase could be stabilised temporary by macrophytes, stabilisation of good water quality requires continuous regulation of fish community. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Two mesocosm experiments investigating the control of summer phytoplankton growth in a small shallow lake

1. Mesocosm experiments were carried out to examine the relative importance of top down (fish predation) and bottom up (nutrient addition) controls on phytoplankton abundance in a small shallow lake, Little Mere, U.K., in 1998 and 1999. These experiments were part of a series at six sites across Europe. 2. In the 1998 experiment, top‐down processes (through grazing of large Cladocera) were important in determining phytoplankton biomass. The lack of plant refugia for zooplankton was probably important in causing an increasing chlorophyll a concentration even at intermediate fish density. Little Mere normally has abundant macrophytes but they failed to develop substantially during both years. Bottom‐up control was not important in 1998, most probably because of high background nutrient concentrations, as a result of nutrient release from the sediments. 3. In 1999 neither top‐down nor bottom‐up processes were significant in determining phytoplankton biomass. Large cladoceran grazers were absent even in the fish‐free enclosures, probably because dominance of cyanobacteria and high phytoplankton biomass made feeding conditions unsuitable. As in 1998, bottom‐up control of phytoplankton was not important, owing to background nutrient concentrations that were even higher in 1999 than in 1998, perhaps because of the warmer, sunnier weather. 4. The differing outcomes of the two experiments in the same lake with similar experimental designs highlight the importance of starting conditions. These conditions in turn depended on overall weather conditions prior to the experiments.