Temperature regulation of marine heterotrophic prokaryotes increases latitudinally as a breach between bottom‐up and top‐down controls

Planktonic heterotrophic prokaryotes make up the largest living biomass and process most organic matter in the ocean. Determining when and where the biomass and activity of heterotrophic prokaryotes are controlled by resource availability (bottom‐up), predation and viral lysis (top‐down) or temperature will help in future carbon cycling predictions. We conducted an extensive survey across subtropical and tropical waters of the Atlantic, Indian and Pacific Oceans during the Malaspina 2010 Global Circumnavigation Expedition and assessed indices for these three types of controls at 109 stations (mostly from the surface to 4,000 m depth). Temperature control was approached by the apparent activation energy in eV (ranging from 0.46 to 3.41), bottom‐up control by the slope of the log‐log relationship between biomass and production rate (ranging from ?0.12 to 1.09) and top‐down control by an index that considers the relative abundances of heterotrophic nanoflagellates and viruses (ranging from 0.82 to 4.83). We conclude that temperature becomes dominant (i.e. activation energy >1.5 eV) within a narrow window of intermediate values of bottom‐up (0.3–0.6) and top‐down 0.8–1.2) controls. A pervasive latitudinal pattern of decreasing temperature regulation towards the Equator, regardless of the oceanic basin, suggests that the impact of global warming on marine microbes and their biogeochemical function will be more intense at higher latitudes. Our analysis predicts that 1°C ocean warming will result in increased biomass of heterotrophic prokaryoplankton only in waters with <26°C of mean annual surface temperature.     

State of the science in reconciling top‐down and bottom‐up approaches for terrestrial CO2 budget

Robust estimates of CO₂ budget, CO₂ exchanged between the atmosphere and terrestrial biosphere, are necessary to better understand the role of the terrestrial biosphere in mitigating anthropogenic CO₂ emissions. Over the past decade, this field of research has advanced through understanding of the differences and similarities of two fundamentally different approaches: “top‐down” atmospheric inversions and “bottom‐up” biosphere models. Since the first studies were undertaken, these approaches have shown an increasing level of agreement, but disagreements in some regions still persist, in part because they do not estimate the same quantity of atmosphere–biosphere CO₂ exchange. Here, we conducted a thorough comparison of CO₂ budgets at multiple scales and from multiple methods to assess the current state of the science in estimating CO₂ budgets. Our set of atmospheric inversions and biosphere models, which were adjusted for a consistent flux definition, showed a high level of agreement for global and hemispheric CO₂ budgets in the 2000s. Regionally, improved agreement in CO₂ budgets was notable for North America and Southeast Asia. However, large gaps between the two methods remained in East Asia and South America. In other regions, Europe, boreal Asia, Africa, South Asia, and Oceania, it was difficult to determine whether those regions act as a net sink or source because of the large spread in estimates from atmospheric inversions. These results highlight two research directions to improve the robustness of CO₂ budgets: (a) to increase representation of processes in biosphere models that could contribute to fill the budget gaps, such as forest regrowth and forest degradation; and (b) to reduce sink–source compensation between regions (dipoles) in atmospheric inversion so that their estimates become more comparable. Advancements on both research areas will increase the level of agreement between the top‐down and bottom‐up approaches and yield more robust knowledge of regional CO₂ budgets.     

Priority effects are interactively regulated by top‐down and bottom‐up forces: evidence from wood decomposer communities

Both top‐down (grazing) and bottom‐up (resource availability) forces can determine the strength of priority effects, or the effects of species arrival history on the structure and function of ecological communities, but their combined influences remain unresolved. To test for such influences, we assembled experimental communities of wood‐decomposing fungi using a factorial manipulation of fungivore (Folsomia candida) presence, nitrogen availability, and fungal assembly history. We found interactive effects of all three factors on fungal species composition and wood decomposition 1 year after the fungi were introduced. The strength of priority effects on community structure was affected primarily by nitrogen availability, whereas the strength of priority effects on decomposition rate was interactively regulated by nitrogen and fungivores. These results demonstrate that top‐down and bottom‐up forces jointly determine how strongly assembly history affects community structure and function.     

Timing of disturbance,top‐down,and bottom‐up driving on early algal succession patterns in a tropical intertidal community

Herbivory and nutrient enrichment are major drivers of the dynamics of algal communities. However, their effects on algal abundance are under the influence of seasons. This study investigated the effects of herbivory and nutrient enrichment on early algal succession patterns using cages (uncaged and fully caged treatments) and two nutrient levels (ambient and enriched concentrations). To determine seasonal influences, experiment plots on dead coral patches were cleared during both dry and rainy season. Of the 17 algal species recruited in the experiment plots, three were dominant: Ulva paradoxa C. Agardh, Padina in the Vaughaniella stage, and Polysiphonia sphaerocarpa Børgesen. In this succession process, U. paradoxa was the earliest colonizer and occupied the cleared plots within the first month after clearing with the highest percentage of 83.33 ± 1.67% to 88.33 ± 9.28%. Then, it was replaced by the late successional algae, Padina in the Vaughaniella stage, and P. sphaerocarpa. The effects of herbivory and nutrient enrichment on algal abundance varied across algal functional groups and seasons. During the dry season, neither herbivory nor nutrient enrichment affected Ulva cover but during the rainy season, Ulva cover was influenced by nutrient enrichment. However, the abundance of algae in this early stage was not apparently affected by either herbivory or nutrient enrichment. Our results indicated that the timing of disturbance strongly influenced the algal abundance and successional patterns in this tropical intertidal community.     

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.     

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.     

Differential responses of size‐based functional groups to bottom–up and top–down perturbations in pelagic food webs: a meta‐analysis

We performed a meta‐analysis of 31 lake mesocosm experiments to investigate differences in the responses of pelagic food chains and food webs to nutrient enrichment and fish presence. Trophic levels were divided into size‐based functional groups (phytoplankton into highly edible and poorly edible algae, and zooplankton into small herbivores, large herbivores and omnivorous zooplankton) in the food webs. Our meta‐analysis shows that 1) nutrient enrichment has a positive effect on phytoplankton and zooplankton, while fish presence has a positive effect on phytoplankton and a negative effect on zooplankton in the food chains; 2) nutrient enrichment has a positive effect on highly edible algae and small herbivores, but no effect on poorly edible algae, large herbivores and omnivorous zooplankton in the food webs. Planktivorous fish have a positive effect on highly edible algae and small herbivores, a negative effect on large herbivores and omnivorous zooplankton, and no effect on poorly edible algae. Our meta‐analysis confirms that nutrient enrichment and planktivorous fish affect functional groups differentially within trophic levels, revealing important changes in the functioning of food webs. The analysis of fish effects shows the well‐described trophic cascade in the food chain and reveals two trophic cascades in the food web: one transmitted by large herbivores that benefit highly edible phytoplankton, and one transmitted by omnivorous zooplankton that benefit small herbivores. Comparison between the responses of food webs and simple food chains also shows consistent biomass compensation between functional groups within trophic levels.     

Bottom‐up and top‐down effects of tree species diversity on leaf insect herbivory

The diversity of plant neighbors commonly results in direct, bottom‐up effects on herbivore ability to locate their host, and in indirect effects on herbivores involving changes in plant traits and a top‐down control by their enemies. Yet, the relative contribution of bottom‐up and top‐down forces remains poorly understood. We also lack knowledge on the effect of abiotic constraints such as summer drought on the strength and direction of these effects. We measured leaf damage on pedunculate oak (Quercus robur), alone or associated with birch, pine or both in a long‐term tree diversity experiment (ORPHEE), where half of the plots were irrigated while the other half remained without irrigation and received only rainfall. We tested three mechanisms likely to explain the effects of oak neighbors on herbivory: (1) Direct bottom‐up effects of heterospecific neighbors on oak accessibility to herbivores, (2) indirect bottom‐up effects of neighbors on the expression of leaf traits, and (3) top‐down control of herbivores by predators. Insect herbivory increased during the growth season but was independent of neighbor identity and irrigation. Specific leaf area, leaf toughness, and thickness varied with neighbor identity while leaf dry matter content or C:N ratio did not. When summarized in a principal component analysis (PCA), neighbor identity explained 87% of variability in leaf traits. PCA axes partially predicted herbivory. Despite greater rates of attack on dummy caterpillars in irrigated plots, avian predation, and insect herbivory remained unrelated. Our study suggests that neighbor identity can indirectly influence insect herbivory in mixed forests by modifying leaf traits. However, we found only partial evidence for these trait‐mediated effects and suggest that more attention should be paid to some unmeasured plant traits such as secondary metabolites, including volatile organic compounds, to better anticipate the effects of climate change on plant‐insect interactions in the future.     

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.     

Top‐down and bottom‐up control of litter decomposers in streams

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Ocean acidification alters zooplankton communities and increases top‐down pressure of a cubozoan predator

The composition of local ecological communities is determined by the members of the regional community that are able to survive the abiotic and biotic conditions of a local ecosystem. Anthropogenic activities since the industrial revolution have increased atmospheric CO₂ concentrations, which have in turn decreased ocean pH and altered carbonate ion concentrations: so called ocean acidification (OA). Single‐species experiments have shown how OA can dramatically affect zooplankton development, physiology and skeletal mineralization status, potentially reducing their defensive function and altering their predatory and antipredatory behaviors. This means that increased OA may indirectly alter the biotic conditions by modifying trophic interactions. We investigated how OA affects the impact of a cubozoan predator on their zooplankton prey, predominantly Copepoda, Pleocyemata, Dendrobranchiata, and Amphipoda. Experimental conditions were set at either current (pCO₂ 370 μatm) or end‐of‐the‐century OA (pCO₂ 1,100 μatm) scenarios, crossed in an orthogonal experimental design with the presence/absence of the cubozoan predator Carybdea rastoni. The combined effects of exposure to OA and predation by C. rastoni caused greater shifts in community structure, and greater reductions in the abundance of key taxa than would be predicted from combining the effect of each stressor in isolation. Specifically, we show that in the combined presence of OA and a cubozoan predator, populations of the most abundant member of the zooplankton community (calanoid copepods) were reduced 27% more than it would be predicted based on the effects of these stressors in isolation, suggesting that OA increases the susceptibility of plankton to predation. Our results indicate that the ecological consequences of OA may be greater than predicted from single‐species experiments, and highlight the need to understand future marine global change from a community perspective.     

Seasonal shifts in the importance of bottom–up and top–down factors on stream periphyton community structure

We examined the importance of temporal variability in top–down and bottom–up effects on the accumulation of stream periphyton, which are complex associations of autotrophic and heterotrophic microorganisms. Periphyton contributes to primary production and nutrient cycling and serves as a food resource for herbivores (grazers). Periphyton growth is often limited by the availability of nitrogen and phosphorus, and biomass can be controlled by grazers. In this study we experimentally manipulated nutrients and grazers simultaneously to determine the relative contribution of bottom–up and top–down controls on periphyton over time. We used nutrient diffusing substrates to regulate nutrient concentrations and an underwater electric field to exclude grazing insects in three sequential 16–17 day experiments from August to October in montane Colorado, USA. We measured algal biomass, periphyton organic mass, and algal community composition in each experiment and determined densities of streambed insect species, including grazers. Phosphorus was the primary limiting nutrient for algal biomass, but it did not influence periphyton organic mass across all experiments. Effects of nutrient additions on algal biomass and community composition decreased between August and October. Grazed substrates supported reduced periphyton biomass only in the first experiment, corresponding to high benthic abundances of a dominant mayfly grazer (Rhithrogena spp.). Grazed substrates in the first experiment also showed altered algal community composition with reduced diatom relative abundances, presumably in response to selective grazing. We showed that top–down grazing effects were strongest in late summer when grazers were abundant. The effects of phosphorus additions on algal biomass likely decreased over time because temperature became more limiting to growth than nutrients, and because reduced current velocity decreased nutrient uptake rates. These results suggest that investigators should proceed with caution when extending findings based on short‐term experiments. Furthermore, these results support the need for additional seasonal‐scale field research in stream ecology.     

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.     

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.