Cascading effects of climate change on plankton community structure

Plankton communities account for at least half of global primary production and play a key role in the global carbon cycle. Warming and acidification may alter the interaction chains in these communities from the bottom and top of the food web. Yet, the relative importance of these potentially complex interactions has not yet been quantified. Here, we examine the isolated and combined effects of warming, acidification, and reductions in phytoplankton and predator abundances in a series of factorial experiments. We find that warming directly impacts the top of the food web, but that the intermediate trophic groups are more strongly influenced by indirect effects mediated by altered top‐down interactions. Direct manipulations of predator and phytoplankton abundance reveal similar strong top‐down interactions following top predator decline. A meta‐analysis of published experiments further supports the conclusion that warming has stronger direct impacts on the top and bottom of the food web rather than the intermediate trophic groups, with important differences between freshwater and marine plankton communities. Our results reveal that the trophic effect of warming cascading down from the top of the plankton food web is a powerful agent of global change.     

Effects of predation and nutrient enrichment on a food web with edible and inedible prey

1. Trophic heterogeneity, or differences in edibility or resource consumption among species within a single trophic level, is widespread in natural food webs. Here, we test simple food web models that incorporate trophic heterogenity and that make robust predictions regarding food web responses to nutrient enrichment. To test these predictions, we assembled simple food webs consisting of an inedible alga, a mixed assemblage of bacteria, and a protist bacterivore in laboratory microcosms of contrasting nutrient concentrations. 2. Several results were consistent with model predictions. First, increasing nutrient concentration caused an increase in the abundance of the inedible alga, but only in the presence of the bacterivore. Secondly, nutrient enrichment increased the abundance of bacteria, but only in the absence of their bacterivore. Last, nutrient enrichment had no effect on abundance of the bacterivore. 3. Two results were not consistent with model predictions. First, at low nutrient concentrations, the presence of the bacterivore increased the abundance of bacteria. Secondly, although the abundances of the bacterivore and bacteria were positively correlated, some of the lowest abundances of both occurred in the high nutrient treatment. Thus, while our results were generally consistent with several simple food web models, additional explanations are required for selected food web responses.     

Trophic cascades and trophic trickles in pelagic food webs

Prey-dependent models, with the predation rate (per predator) a function of prey numbers alone, predict the existence of a trophic cascade. In a trophic cascade, the addition of a top predator to a two-level food chain to make a three-level food chain will lead to increases in the population size of the primary producers, and the addition of nutrients to three-level chains will lead to increases in the population numbers at only the first and third trophic levels. In contrast, ratio-dependent models, with the predation rate (per predator) dependent on the ratio of predator numbers to prey, predict that additions of top predators will not increase the population sizes of the primary producers, and that the addition of nutrients to a three-level food chain will lead to increases in population numbers at all trophic levels. Surprisingly, recent meta-analyses show that freshwater pelagic food web patterns match neither prey-dependent models (in pelagic webs, ''prey'' are phytoplankton, and ''predators'' are zooplankton), nor ratio-dependent models. In this paper we use a modification of the prey-dependent model, incorporating strong interference within the zooplankton trophic level, that does yield patterns matching those found in nature. This zooplankton interference model corresponds to a more reticulate food web than in the linear, prey-dependent model, which lacks zooplankton interference. We thus reconcile data with a new model, and make the testable prediction that the strength of trophic cascades will depend on the degree of heterogeneity in the zooplankton level of the food chain.     

Effect of parity on productivity and sustainability of Lotka–Volterra food chains

Hairston, Slobodkin, and Smith conjectured that top down forces act on food chains, which opposed the previously accepted theory that bottom up forces exclusively dictate the dynamics of populations. We model food chains using the Lotka–Volterra predation model and derive sustainability constants which determine which species will persist or go extinct. Further, we show that the productivity of a sustainable food chain with even trophic levels is predator regulated, or top down, while a sustainable food chain with odd trophic levels is resource limited, which is bottom up, which is consistent with current ecological theory.     

Most soil trophic guilds increase plant growth: a meta‐analytical review

Trophic cascades are important drivers of plant and animal abundances in aquatic and aboveground systems, but in soils trophic cascades have been thought to be of limited importance due to omnivory and other factors. Here we use a meta‐analysis of 215 studies with 1526 experiments that measured plant growth responses to additions or removals of soil organisms to test how different soil trophic levels affect plant growth. Consistent with the trophic cascade hypothesis, we found that herbivores and plant pathogens (henceforth pests) decreased plant growth and that predators of pests increased plant growth. The magnitude of this trophic cascade was similar to that reported for aboveground systems. In contrast, we did not find evidence for trophic cascades in decomposer‐ and symbiont‐based (henceforth mutualist) food chains. In these food chains, mutualists increased plant growth and predators of mutualists also increased plant growth, presumably by increasing nutrient cycling rates. Therefore, mutualists, predators of mutualists and predators of pests all increased plant growth. Further, experiments that added multiple organisms from different trophic levels also increased plant growth. As a result, across the dataset, soil organisms increased plant growth 29% and non‐pest soil organisms increased plant growth 46%. Omnivory has traditionally been thought to confound soil trophic dynamics, but here we suggest that omnivory allows for a simplified perspective of soil food webs – one in which most soil organisms increase plant growth by preying on pests or increasing nutrient cycling rates. An implication of this perspective is that processes that decrease soil organism abundance (e.g. soil tillage) are likely to decrease aboveground productivity. Synthesis Soil foodwebs have resisted generalizations due to their diversity and interconnectedness. Here we use results from a meta‐analysis to inform a simplified perspective of soil foodwebs: one in which most soil trophic guilds increase plant growth. Our review also includes the first widespread support for the presence of trophic cascades in soils.     

Ratio-dependent response of a temperate Australian estuarine system to sustained nitrogen loading

Classical resource- and the less studied ratio-dependent models of predator–prey relationships provide divergent predictions as to the sustained ecological effects of bottom-up forcing. While resource-dependent models, which consider only instantaneous prey density in modelling predator responses, predict community responses that are dependent on the number of trophic levels in a system, ratio-dependent models, which consider the number of prey per consumer, predict proportional increase in each level irrespective of chain length. The two models are only subtly different for systems with two or three trophic levels but in the case of four trophic levels, predict opposite effects of enrichment on primary producers. Despite the poor discriminatory power of tests of the models in systems with two or three trophic levels, field tests in estuarine and marine systems with four trophic levels have been notably absent. Sampling of phytoplankton, macroinvertebrates, invertebrate-feeding fishes, piscivorous fishes in Kooloonbung Creek, Hastings River estuary, eastern Australia, subject to over 20 years of sewage discharge, revealed increased abundances in all four trophic levels at the disturbed location relative to control sites. Increased abundance of phytoplankton at the disturbed site was counter to the predictions of resource-dependent models, which posit a reduction in the first trophic level in response to enrichment. By contrast, the increase in abundance of this first trophic level and the proportionality of increases in abundances of each of the four trophic groups to nitrogen loading provided strong support for ratio dependency. This first evidence of ratio dependence in an estuarine system with four trophic levels not only demonstrates the applicability of ecological theory which seeks to simplify the complexity of systems, but has implications for management. Although large nutrient inputs frequently induce mortality of invertebrates and fish, we have shown that smaller inputs may in fact enhance biomass of all trophic levels.     

Understanding patterns and processes in models of trophic cascades

Climate fluctuations and human exploitation are causing global changes in nutrient enrichment of terrestrial and aquatic ecosystems and declining abundances of apex predators. The resulting trophic cascades have had profound effects on food webs, leading to significant economic and societal consequences. However, the strength of cascades–that is the extent to which a disturbance is diminished as it propagates through a food web–varies widely between ecosystems, and there is no formal theory as to why this should be so. Some food chain models reproduce cascade effects seen in nature, but to what extent is this dependent on their formulation? We show that inclusion of processes represented mathematically as density‐dependent regulation of either consumer uptake or mortality rates is necessary for the generation of realistic ‘top‐down’ cascades in simple food chain models. Realistically modelled ‘bottom‐up’ cascades, caused by changing nutrient input, are also dependent on the inclusion of density dependence, but especially on mortality regulation as a caricature of, e.g. disease and parasite dynamics or intraguild predation. We show that our conclusions, based on simple food chains, transfer to a more complex marine food web model in which cascades are induced by varying river nutrient inputs or fish harvesting rates.     

Effects of Multi-chain Omnivory on the Strength of Trophic Control in Lakes

Omnivory has been implicated in both diffusing and intensifying the effects of consumer control in food chains. Some have postulated that the strong, community level, top-down control apparent in lakes is not expressed in terrestrial systems because terrestrial food webs are reticulate, with high degrees of omnivory and diverse plant communities. In contrast, lake food webs are depicted as simple linear chains based on phytoplankton-derived energy. Here, we explore the dynamic implications of recent evidence showing that attached algal (periphyton) carbon contributes substantially to lake primary and secondary productivity, including fish production. Periphyton production represents a cryptic energy source in oligotrophic and mesotrophic lakes that is overlooked by previous theoretical treatment of trophic control in lakes. Literature data demonstrate that many fish are multi-chain omnivores, exploiting food chains based on both littoral and pelagic primary producers. Using consumer-resource models, we examine how multiple food chains affect fourth-level trophic control across nutrient gradients in lakes. The models predict that the stabilizing effects of linked food chains are strongest in lakes where both phytoplankton and periphyton contribute substantially to production of higher trophic levels. This stabilization enables a strong and persistent top down control on the pelagic food chain in mesotrophic lakes. The extension of classical trophic cascade theory to incorporate more complex food web structures driven by multi-chain predators provides a conceptual framework for analysis of reticulate food webs in ecosystems.     

Top‐down and bottom‐up diversity cascades in detrital vs. living food webs

Apex predators and plant resources are both critical for maintaining diversity in biotic communities, but the indirect (‘cascading’) effects of top‐down and bottom‐up forces on diversity at different trophic levels are not well resolved in terrestrial systems. Manipulations of predators or resources can cause direct changes of diversity at one trophic level, which in turn can affect diversity at other trophic levels. The indirect diversity effects of resource and consumer variation should be strongest in aquatic systems, moderate in terrestrial systems, and weakest in decomposer food webs. We measured effects of top predators and plant resources on the diversity of endophytic animals in an understorey shrub Piper cenocladum (Piperaceae). Predators and resource availability had significant direct and indirect effects on the diversity of the endophytic animal community, but the effects were not interactive, nor were they consistent between living vs. detrital food webs. The addition of fourth trophic level beetle predators increased diversity of consumers supported by living plant tissue, whereas balanced plant resources (light and nutrients) increased the diversity of primary through tertiary consumers in the detrital resources food web. These results support the hypotheses that top‐down and bottom‐up diversity cascades occur in terrestrial systems, and that diversity is affected by different factors in living vs. detrital food webs.     

Upwelling-like bottom intrusion enhances the pelagic–benthic coupling by a fish predator in a coastal food web

Upwelling regions where nutrients are transported from deep to surface waters are among the most productive in the oceans. Although it is well known that the upwelling affects fishery production through bottom-up trophic cascading, it remains unexplored how temporal variation in its intensity alters overall trophic energy flows within a focal food web. In the present study, we demonstrate that inter-annual variation in the intensity of upwelling-like bottom intrusion alters food web properties in coastal waters of the Uwa Sea by focusing on the levels of δ¹³C and δ¹⁵N for a demersal fish predator, Acropoma japonicum. This approach integrates information on prey–predator interactions. In the season following a stratification period when pelagic productivity is limited by nutrient availability, A. japonicum showed lower levels of δ¹³C in years with high bottom intrusion intensity than in those with low intensity. One possible cause for this isotopic depletion is that the bottom intrusion-induced nutrient supply enhances pelagic productivity and consequently facilitates a foraging shift by A. japonicum from ordinary benthic prey to supplementary pelagic prey with a lower δ¹³C. In conclusion, the increased intensity of bottom intrusion results in coupling of two major trophic energy flows, pelagic and benthic food chains, through the demersal predator’s foraging shift.     

Biodiversity and ecosystem functioning in food webs: the vertical diversity hypothesis

One challenge in merging community and ecosystem ecology is to integrate the complexity of natural multitrophic communities into concepts of ecosystem functioning. Here, we combine food‐web and allometry theories to demonstrate that primary production, as measured by the total nutrient uptake of the multitrophic community, is determined by vertical diversity (i.e. food web's maximum trophic level) and structure (i.e. distributions of species and their abundances and metabolic rates across trophic levels). In natural ecosystems, the community size distribution determines all these vertical patterns and thus the total nutrient uptake. Our model suggests a vertical diversity hypothesis (VDH) for ecosystem functioning in complex food webs. It predicts that, under a given nutrient supply, the total nutrient uptake increases exponentially with the maximum trophic level in the food web and it increases with its maximum body size according to a power law. The VDH highlights the effect of top–down regulation on plant nutrient uptake, which complements traditional paradigms that emphasised the bottom–up effect of nutrient supply on vertical diversity. We conclude that the VDH contributes to a synthetic framework for understanding the relationship between vertical diversity and ecosystem functioning in food webs and predicting the impacts of global changes on multitrophic ecosystems.     

氮磷输入对过度放牧退化草原土壤线虫群落的影响

线虫是陆地生态系统中数量最多的一类无脊椎后生动物,在土壤碎屑食物网中占据多个营养级,并在能量流动和养分循环中发挥重要的生态功能.土壤线虫的群落结构和多样性水平可以反映生态系统功能的变化.多年来,过度放牧等不合理的开发利用导致我国大面积的草原处在不同程度的退化之中,外源性养分输入是维持其养分平衡进而恢复其生态和生产功能的重要措施.本研究以内蒙古锡林郭勒退化草原为对象,就氮磷输入对土壤线虫丰度和群落结构的单独效应和交互效应进行了研究.本研究获得线虫38属,其中垫咽属、滑刃属、索努斯属和盾垫属是该退化草原线虫群落优势属.氮输入降低了土壤线虫总丰度、捕食杂食线虫丰度以及植物寄生线虫丰度,而磷输入提高了土壤线虫总丰度、食真菌线虫丰度、捕食杂食类线虫丰度以及植物寄生线虫丰度,氮输入抑制了磷输入对线虫总丰度、捕食杂食线虫和植物寄生线虫丰度的促进作用.氮磷输入对线虫多样性无影响,可能与不同养分输入下稳定的植物群落多样性有关.氮输入显著提升了退化草原线虫成熟度指数,降低了植物寄生线虫成熟度指数(PPI),并且能缓解磷输入对PPI和瓦斯乐斯卡指数的负面作用,表明氮输入提高退化草地土壤健康程度,促进线虫群落和食物网的稳定性.上述研究结果有助于从土壤生物学角度理解氮磷输入对退化草地恢复的影响机理.     

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.     

Allochthonous inputs: integrating population changes and food-web dynamics

Most ecosystems are recipients of allochthonous materials that enhance in situ productivity. Recent theoretical and empirical studies suggest that low to moderate inputs can stabilize food webs. However, depending on the trophic levels that use the resource, food webs can become unstable as inputs increase. Where large amounts of agricultural resources are transferred to natural habitats, trophic dynamics change: trophic cascades can occur and rare or uncommon species can become invasive. Rates of change in species abundances can also be amplified by the effects of changes in legislation and management practices on subsidized consumers.     

Bottom-up effects may not reach the top: the influence of ant-aphid interactions on the spread of soil disturbances through trophic chains

Soil disturbances that increase nutrient availability may trigger bottom-up cascading effects along trophic chains. However, the strength and sign of these effects may depend on attributes of the interacting species. Here, we studied the effects of nutrient-rich refuse dumps of the leaf-cutting ant, Acromyrmex lobicornis, on the food chain composed of thistles, aphids, tending ants and aphid natural enemies. Using stable isotopes tracers, we show that the nitrogen accumulated in refuse dumps propagates upward through the studied food chain. Thistles growing on refuse dumps had greater biomass and higher aphid density than those growing in adjacent soil. These modifications did not affect the structure of the tending ant assemblage, but were associated with increased ant activity. In contrast to the expectations under the typical bottom-up cascade effect, the increase in aphid abundance did not positively impact on aphid natural enemies. This pattern may be explained by both an increased activity of tending ants, which defend aphids against their natural enemies, and the low capacity of aphid natural enemies to show numerical or functional responses to increased aphid density. Our results illustrate how biotic interactions and the response capacity of top predators could disrupt bottom-up cascades triggered by disturbances that increase resource availability.     

Carnivore identity mediates the effects of light and nutrients on aquatic food‐chain efficiency

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The dynamics of top-down and bottom-up effects in food webs of varying prey diversity, composition, and productivity

Prey diversity is thought to mediate the strength of top-down and bottom-up effects, but few experiments directly test this hypothesis. I assembled food webs of bacteria and bacterivorous protist prey in laboratory microcosms with all combinations of five productivity levels, two top predator treatments (present or absent), and three prey compositions. Depauperate food chains contained one of two edible prey species, while more diverse food webs contained both edible prey species plus two additional less-edible/inedible prey. Equilibrium theory predicts that prey diversity should weaken the top-down and bottom-up effects on trophic level biomasses, due to density compensation among prey species. Top-down effects should increase with productivity in food chains, but decrease with productivity in food webs. Results revealed highly dynamic top-down effects, the strength of which varied more over time than among treatments. Further, top-down effects did not merely vary in absolute strength over time, but also in relative strength across different prey compositions and productivity levels. It might be expected that equilibrium models would qualitatively reproduce time-averaged results. However, time-averaged data largely failed to support equilibrium predictions. This failure may reflect strong temporal variability in treatment effects combined with nonlinear density dependence of species' per-capita growth rates. Strong temporal variability in the strength of top-down effects has not previously been demonstrated, but likely is common in nature as well.     

A damped precipitation‐driven,bottom‐up model for deer mouse population abundance in the northwestern United States

Small‐mammal population densities can be regulated by bottom‐up (food availability) and top‐down (predation) forces. In 1993, an El Niño Southern Oscillation event was followed by a cluster of human hantavirus with pulmonary syndrome in the southwestern United States. An upward trophic cascade hypothesis was proposed as an explanation for the outbreak: Increased plant productivity as a consequence of El Niño precipitations led to an unusual increase in distribution and abundance of deer mice (Peromyscus maniculatus ; reservoir host of Sin Nombre virus). Could such drastic events occur in mesic habitats, where plant productivity in response to climate conditions is likely to be much less dramatic? In this work, we investigate to what extent deer mouse populations follow a precipitation‐driven, bottom‐up model in central and western Montana and discuss important conditions for such a model to be possible. We found positive correlations between deer mouse abundance and on‐the‐ground measured plant productivity with a several‐month lag in three of six study sites. This effect was weaker when deer mouse populations were more abundant, indicating density‐dependent effects. Dispersal resulting from territoriality may be important in attenuating local density increments in spite of high food availability. In addition, there is evidence that population abundance in the study area could respond to other abiotic factors. In particular, precipitation in the form of snow may reduce deer mice survival, thus compensating the benefits of improved plant productivity. Deer mouse populations in Montana study sites follow complex dynamics determined by multiple limiting factors, leading to a damped precipitation‐driven bottom‐up regulation. This prevents dramatic changes in rodent abundances after sudden increments of food availability, such as those observed in other regions.