互利关系和植物多样性对节肢动物群落不同营养级的影响路径和强度

食物网中的上行效应和下行效应对于群落的动态和生态系统功能有十分重要的影响,旨在探讨互利关系和植物多样性对节肢动物群落中食物网不同营养级之间的影响。通过随机裂区试验方法,分别设置了3种蚂蚁-紫胶虫互利关系处理(有互利关系、无互利关系和自然对照)以及3种植物多样性处理(单一种植、2树种混植和3树种混植),于2016年8月和9月分两次用手捡法、网扫法和震落法采集试验地寄主植物上所有的节肢动物,并按照不同营养级将其分类。利用结构方程模型分析方法对不同营养级之间的相互作用的路径和强度进行了比较,结果显示:1)互利关系对捕食者和消费者均有显著的下行作用,有互利关系处理下蚂蚁对捕食者的路径强度要强于自然对照组,互利关系对捕食者的影响要强于对消费者的影响。2)植物多样性会通过影响植物的生物量而对消费者和捕食者产生显著的上行效应影响,这种影响会随着营养级的升高而显著减小。3)消费者主要受植物多样性的上行效应影响,而捕食者主要受互利关系的下行效应影响。有互利关系的食物网结构更加复杂,营养级之间的相互作用更为显著。探讨了以蚂蚁-紫胶虫互利关系为核心作用的紫胶林生态系统中互利关系和植物多样性对节肢动物食物网中...     

Evaluating trophic cascades as drivers of regime shifts in different ocean ecosystems

In ecosystems that are strongly structured by predation, reducing top predator abundance can alter several lower trophic levels—a process known as a trophic cascade. A persistent trophic cascade also fits the definition of a regime shift. Such ‘trophic cascade regime shifts'' have been reported in a few pelagic marine systems—notably the Black Sea, Baltic Sea and eastern Scotian Shelf—raising the question of how common this phenomenon is in the marine environment. We provide a general methodology for distinguishing top-down and bottom-up effects and apply this methodology to time series from these three ecosystems. We found evidence for top-down forcing in the Black Sea due primarily to gelatinous zooplankton. Changes in the Baltic Sea are primarily bottom-up, strongly structured by salinity, but top-down forcing related to changes in cod abundance also shapes the ecosystem. Changes in the eastern Scotian Shelf that were originally attributed to declines in groundfish are better explained by changes in stratification. Our review suggests that trophic cascade regime shifts are rare in open ocean ecosystems and that their likelihood increases as the residence time of water in the system increases. Our work challenges the assumption that negative correlation between consecutive trophic levels implies top-down forcing.     

Complex wasp-waist regulation of pelagic ecosystems in the Pacific Ocean

‘Wasp-waist’ control of marine ecosystems is driven by a combination of top-down and bottom-up forcing by a few abundant short-lived species occupying intermediate trophic levels that form a narrow ‘waist’ through which energy flow from low to high trophic levels is controlled. It has been assumed that wasp-waist control occurs primarily in highly productive and species-poor systems (e.g. upwelling regions). Two large, species-rich, pelagic ecosystems in the relatively oligotrophic eastern and western Pacific Ocean also show wasp-waist-like structure, in that short-lived and fast-growing cephalopods and fishes at intermediate trophic levels comprise the vast majority of the biomass. Possible forcing dynamics of these systems were examined using ecosystem models by altering the biomass of phytoplankton (bottom-up forcing), large pelagic predators (top-down forcing), and intermediate ‘wasp-waist’ functional groups independently and observing how these changes propagated throughout the ecosystem. The largest effects were seen when altering the biomass of mid trophic-level epipelagic and mesopelagic fishes, where dramatic trophic cascades occurred both upward and downward in the system. We conclude that the high productivity and standing biomass of animals at intermediate trophic levels has a strong top-down influence on the abundance of primary producers. Furthermore, their importance as prey for large predators results in bottom-up controls on populations at higher trophic levels. We show that these tropical pelagic ecosystems possess a complex structure whereby several waist groups and alternate trophic pathways from primary producers to apex predators can cause unpredictable effects when the biomasses of particular functional groups are altered. Such models highlight the possible structuring mechanisms in pelagic systems, which have implications for fisheries that exploit these wasp-waist groups, such as squid fisheries, as well as for fisheries of top predators such as tunas and billfishes that prey upon wasp-waist species.     

A test of three alternative pathways for consumer regulation of primary productivity

The pathways linking consumer effects to primary productivity (PPR) are likely to vary among taxa because of species-specific trophic and functional differences. Thus, it is necessary to understand the dynamics of consumer–PPR interactions so that effects of species loss on ecosystem function can be addressed from a mechanistic approach. In this study, I used three fish taxa (orangethroat darter, Etheostoma spectabile; western mosquitofish, Gambusia affinis; and bullhead minnow, Pimephales vigilax) as model consumers with different trophic and functional characteristics to test alternative mechanisms for consumer regulation of PPR (i.e., trophic cascade, terrestrial nutrient translocation, and sedimentary nutrient translocation). Experiments were conducted in stream mesocosms fitted with a combination of fish and terrestrial insect barriers to address relative importance of consumer-driven top-down and bottom-up control of PPR. A predatory invertivore, orangethroat darter, increased PPR through an apparent trophic cascade by localized reduction of benthic grazing invertebrate densities (i.e., top-down). A surface feeding insectivore, western mosquitofish, consumed terrestrial insects on the stream surface, increasing PPR by enhancing allochthonous nutrients in the mesocosms (i.e., bottom-up). A benthic omnivore, bullhead minnow, consumed benthic food items, resulting in increased PPR by enhancing availability of autochthonous nutrients via translocation of sedimentary nutrients (i.e., bottom-up). However, under specific environmental contexts, this species also consumed terrestrial invertebrates, potentially affecting PPR through terrestrial nutrient translocation as well. In this study, the trophic and functional characteristics of different species resulted in alternative pathways that increased PPR, suggesting that in natural ecosystems multiple consumer-driven pathways may be influencing PPR simultaneously and could potentially be important for temporal persistence of ecosystem function in changing environments.     

The ups and downs of trophic control in continental shelf ecosystems

Traditionally, marine ecosystem structure was thought to be determined by phytoplankton dynamics. However, an integrated view on the relative roles of top-down (consumer-driven) and bottom-up (resource-driven) forcing in large-scale, exploited marine ecosystems is emerging. Long time series of scientific survey data, underpinning the management of commercially exploited species such as cod, are being used to diagnose mechanisms that could affect the composition and relative abundance of species in marine food webs. By assembling published data from studies in exploited North Atlantic ecosystems, we found pronounced geographical variation in top-down and bottom-up trophic forcing. The data suggest that ecosystem susceptibility to top-down control and their resiliency to exploitation are related to species richness and oceanic temperature conditions. Such knowledge could be used to produce ecosystem guidelines to regulate and manage fisheries in a sustainable fashion.     

Nonadditive interactions between trophic levels bias the appraisal of the strength of mortality factors

The role of bottom-up and top-down factors in determining the abundance of organisms is still often evaluated in terms of the nominal (i.e., apparent) cause of death. To determine whether estimates of mortality can be influenced by nonadditive multitrophic interactions, we generated both marginal (i.e., the entirety of mortality attributable to a given factor, including its obscured fraction) and nominal estimates of the trophic forces acting on an herbivore. This was accomplished by comparing mortality rates of the willow leaf beetle Phratora vulgatissima on willows affected or relatively unaffected by local growing conditions, in the presence or absence of natural enemies. Marginal estimates indicated that bottom-up and top-down mortality factors interacted in a synergistic or compensatory way, with the nature of the interaction varying in response to the time elapsed since the last harvest of willow plantations. In contrast, nominal estimates could not discriminate synergistic and compensatory interactions from top-down influence. Combined with recent evidence of compensatory mortality between trophic forces in another system, this study suggests that nonadditive effects between bottom-up and top-down mortality factors may be common, offering an explanation through which the contrasting evidences previously presented by proponents of the bottom-up and top-down views can be understood.     

Trophic exploitation in grassland food chains: simple models and a field experiment

This study provides insight into the importance of top carnivores (top-down control) and nutrient inputs (bottom-up control) in structuring food chains in a terrestrial grassland system. Qualitative predictions about food chain structure are generated using 4 simple models, each differing in assumptions about some key component in the population dynamics of the herbivore trophic level. The four model systems can be classified broadly into two groups (1) those that assume plant resource intake by herbivores is limited by search rate and handling time as described by classic Lotka-Volterra models; and (2) those that assume plant resource intake by herbivores is limited externally by the supply rate of resources as described by alternatives to Lotka-Volterra formulations. The first class of models tends to ascribe greater importance to top-down control of food chain structure whereas the second class places greater weight on bottom-up control. I evaluated the model predictions using experimentally assembled grassland food chains in which I manipulated nutrient inputs and carnivore (wolf spider) abundance to determine the degree of top-down and bottom-up control of grassland plants and herbivores (grasshoppers). The experimental results were most consistent with predictions of the second class of models implying a predominance of bottom-up control of food chain structure.     

Combined Fishing and Climate Forcing in the Southern Benguela Upwelling Ecosystem: An End-to-End Modelling Approach Reveals Dampened Effects

The effects of climate and fishing on marine ecosystems have usually been studied separately, but their interactions make ecosystem dynamics difficult to understand and predict. Of particular interest to management, the potential synergism or antagonism between fishing pressure and climate forcing is analysed in this paper, using an end-to-end ecosystem model of the southern Benguela ecosystem, built from coupling hydrodynamic, biogeochemical and multispecies fish models (ROMS-N₂P₂Z₂D₂-OSMOSE). Scenarios of different intensities of upwelling-favourable wind stress combined with scenarios of fishing top-predator fish were tested. Analyses of isolated drivers show that the bottom-up effect of the climate forcing propagates up the food chain whereas the top-down effect of fishing cascades down to zooplankton in unfavourable environmental conditions but dampens before it reaches phytoplankton. When considering both climate and fishing drivers together, it appears that top-down control dominates the link between top-predator fish and forage fish, whereas interactions between the lower trophic levels are dominated by bottom-up control. The forage fish functional group appears to be a central component of this ecosystem, being the meeting point of two opposite trophic controls. The set of combined scenarios shows that fishing pressure and upwelling-favourable wind stress have mostly dampened effects on fish populations, compared to predictions from the separate effects of the stressors. Dampened effects result in biomass accumulation at the top predator fish level but a depletion of biomass at the forage fish level. This should draw our attention to the evolution of this functional group, which appears as both structurally important in the trophic functioning of the ecosystem, and very sensitive to climate and fishing pressures. In particular, diagnoses considering fishing pressure only might be more optimistic than those that consider combined effects of fishing and environmental variability.     

Ecosystem alteration modifies the relative strengths of bottom-up and top-down forces in a herbivore population

1. Ecosystem alterations can affect the abundance, distribution and diversity of plants and animals, and thus potentially change the relative strength of bottom-up (the plant resource) and top-down (natural enemies) trophic forces acting on herbivore populations. 2. The hypothesis that alterations of the forest ecosystem associated with precommercial thinning have contributed to the increased severity of outbreaks of Neodiprion abietis (Harris), a sawfly defoliator, through the reduction of trophic forces acting on N. abietis larvae, was tested using exclusion techniques. 3. The relative contributions to N. abietis larval mortality of bottom-up and top-down forces both increased with increasing levels of defoliation and were both reduced by thinning. The reduction of bottom-up and top-down forces caused a 58% mean increase in N. abietis larval survival in thinned compared with untreated stands, which is less than would be expected by the sum of the effects of thinning on each source of mortality. Evidence indicates that the partly compensatory, partly additive nature of the mortality associated with trophic forces in the system under study is responsible for this discrepancy. 4. To our knowledge, this is the first study to show the impact of ecosystem alterations on the balance between bottom-up and top-down forces acting on an eruptive herbivore population along a gradient of host-plant defoliation, and how this can lead to increased outbreak severity. It is stressed that accurate estimates of the relative contributions of bottom-up and top-down forces to mortality cannot be obtained if the additive or compensatory nature of the mortality associated with these trophic forces is overlooked.     

From bottom-up to top-down control of invertebrate herbivores in a retrogressive chronosequence

In the long-term absence of disturbance, ecosystems often enter a decline or retrogressive phase which leads to reductions in primary productivity, plant biomass, nutrient cycling and foliar quality. However, the consequences of ecosystem retrogression for higher trophic levels such as herbivores and predators, are less clear. Using a post-fire forested island-chronosequence across which retrogression occurs, we provide evidence that nutrient availability strongly controls invertebrate herbivore biomass when predators are few, but that there is a switch from bottom-up to top-down control when predators are common. This trophic flip in herbivore control probably arises because invertebrate predators respond to alternative energy channels from the adjacent aquatic matrix, which were independent of terrestrial plant biomass. Our results suggest that effects of nutrient limitation resulting from ecosystem retrogression on trophic cascades are modified by nutrient-independent variation in predator abundance, and this calls for a more holistic approach to trophic ecology to better understand herbivore effects on plant communities.     

Bottom-Up Effects on Biomass Versus Top-Down Effects on Identity: A Multiple-Lake Fish Community Manipulation Experiment

The extent to which ecosystems are regulated by top-down relative to bottom-up control has been a dominant paradigm in ecology for many decades. For lakes, it has been shown that predation by fish is an important determinant of variation in zooplankton and phytoplankton community characteristics. Effects of fish are expected to not only be a function of total fish biomass, but also of functional composition of the fish community. Previous research on the importance of trophic cascades in lakes has largely focused on the role of zooplanktivorous and piscivorous fish. We conducted a large-scale multiple-lake fish community manipulation experiment to test for the effect of differences in fish functional community composition on the trophic structure of lakes. We examine the effect of top-down and bottom-up factors on phytoplankton and zooplankton biomass as well as on their community composition. We put our data in a broader perspective by comparing our results to data of a survey that also included ponds with low fish densities as well as ponds with very high densities of fish. Our results indicate that the overall food web structure under relative high fish densities is primarily structured by bottom-up factors, whereas community characteristics seem to be primarily regulated by top-down factors. Our results suggest a subtle interplay between bottom-up and top-down factors, in which bottom-up factors dominate in determining quantities while top-down effects are important in determining identities of the communities.     

Bottom-up trophic cascades and material transfer in terrestrial food webs

In contrast to top-down trophic cascades, few reviews have appeared of bottom-up trophic cascades. We review the recent development of research on bottom-up cascades in terrestrial food webs, focusing on tritrophic systems consisting of plants, herbivorous insects, and natural enemies, and attempt to integrate bottom-up cascade and material transfer among trophic levels. Bottom-up cascades are frequently reported in various tritrophic systems, and are important to determine community structure, population dynamics, and individual performance of higher trophic levels. In addition, we highlight several features of bottom-up cascades. Accumulation or dilution of plant nutritional and defensive materials by herbivorous insects provides a mechanistic base for several bottom-up cascades. Such a stoichiometric approach has the potential to improve our understanding of bottom-up cascading effects in terrestrial food webs. We suggest a future direction for research by integration of bottom-up cascades and material transfer among trophic levels.     

Relative importance of top-down and bottom-up forces in food webs of Sarracenia pitcher communities at a northern and a southern site

The relative importance of resources (bottom-up forces) and natural enemies (top-down forces) for regulating food web dynamics has been debated, and both forces have been found to be critical for determining food web structure. How the relative importance of top-down and bottom-up forces varies between sites with different abiotic conditions is not well understood. Using the pitcher plant inquiline community as a model system, I examine how the relative importance of top-down and bottom-up effects differs between two disparate sites. Resources (ant carcasses) and top predators (mosquito larvae) were manipulated in two identical 4 × 4 factorial press experiments, conducted at two geographically distant sites (Michigan and Florida) within the range of the purple pitcher plant, Sarracenia purpurea, and the aquatic community that resides in its leaves. Overall, top predators reduced the density of prey populations while additional resources bolstered them, and the relative importance of top-down and bottom-up forces varied between sites and for different trophic levels. Specifically, top-down effects on protozoa were stronger in Florida than in Michigan, while the opposite pattern was found for rotifers. These findings experimentally demonstrate that the strength of predator–prey interactions, even those involving the same species, vary across space. While only two sites are compared in this study, I hypothesize that site differences in temperature, which influences metabolic rate, may be responsible for variation in consumer–resource interactions. These findings warrant further investigation into the specific factors that modify the relative importance of top-down and bottom-up effects.     

Experimental tests of trophic dynamics: towards a more penetrating approach

The complex interactions between primary producers, herbivores, carnivores, and detritivores have resulted in the burgeoning field of trophic dynamics. One important contribution is the Fretwell and Oksanen theory (FO theory). The FO theory proposes that the productivity of the environment determines the length of the trophic chain, which, they suggest, is directly related to whether the system is being controlled by top-down forces (odd numbered length of trophic chain) or bottom-up forces (even numbered length of trophic chain). Recent evidence from experiments by L.H. Fraser and J.P. Grime claims to support the FO theory but the methodology has been criticised by D.C. Moon, P. Stiling and M.V. Cattell for hidden treatments and pseudoreplication. We reject these criticisms and recommend an approach to the study of trophic dynamics involving the aggregation of organisms into functional groups, direct quantitative measurements of trophic processes using field manipulations, inferences based upon the use of field probes and synthesis of ecosystems in closed microcosms. Received: 15 December 1998 / Accepted: 10 December 1998     

Control of arthropod abundance, richness, and composition in a heterogeneous desert city

There is a demand for mechanistic studies to explore underlying drivers behind observed patterns of biodiversity in urban areas. We describe a two-year field experiment in which we manipulated bottom-up (resource availability) and top-down (bird predation) forces on arthropod communities associated with a native plant, Encelia farinosa, across three land-use types—urban, desert remnant, and outlying natural desert—in the Phoenix metropolitan area, Arizona, USA. We monitored the trophic structure, richness, and similarity of the arthropod communities on these manipulated plants over a two-year period. We predicted that (1) increased water resources increase plant productivity, (2) increased productivity increases arthropod abundances, and (3) in the urban habitat, top-down forces are greater than in other habitats and limit arthropod abundances. We also predicted that urban remnant habitats are more similar to urban habitats in terms of arthropod richness and composition. Strong interannual differences due to an unusual cold and dry winter in the first year suppressed plant growth in all but urban habitats, and arthropod abundances in all habitats were severely reduced. In the following year, arthropod abundances in desert and remnant habitats were higher than in urban habitats. Water had positive effects on plant growth and arthropod abundance, but these water effects emerged through complex interactions with habitat type and the presence/absence of cages used to reduce bird predation. Plants grew larger in urban habitats, and phenology also differed between urban and desert habitats. The results from caging suggest that bird predation may not be as important in cities as previously thought, and that arthropods may retard plant growth. As expected, desert communities are strongly bottom-up regulated, but contrary to predictions, we did not find evidence for strong top-down control in the city. Remnant habitats were intermediate between desert and urban habitats in terms of diversity, richness, evenness, arthropod composition and phenology, with urban habitats generally lowest in terms of diversity, richness, and evenness. Our study shows that control of biodiversity is strongly altered in urban areas, influenced by subtle shifts in top-down and bottom-up controls that are often superseded by climatic variations and habitat type.     

Testing hypotheses of trophic level interactions: a boreal forest ecosystem

Models of community organization involve variations of the top-down (predator control) or bottom-up (nutrient limitation) hypotheses. Verbal models, however, can be interpreted in different ways leading to confusion. Therefore, we predict from first principles the range of possible trophic level interactions, and define mathematically the instantaneous effects of experimental perturbations. Some of these interactions are logically and biologically unfeasible. The remaining set of 27 feasible models is based on an initial assumption, for simplicity, of linear interactions between trophic levels. Many more complex and non-linear models are logically feasible but, for parsimony, simple ones are tested first. We use an experiment in the boreal forest of Canada to test predictions of instantaneous changes to trophic levels and distinguish between competing models. Seven different perturbations systematically removed each trophic level or, for some levels, supplemented them. The predictions resulting from the perturbations were concerned with the direction of change in biomass in the other levels. The direct effects of each perturbation produced strong top-down and bottom-up changes in biomass. At both the vegetation and herbivore levels top-down was stronger than bottom-up despite some compensatory growth stimulated by herbivory. The combination of experiments produced results consistent with two-way (reciprocal) interactions at each level. Indirect effects on one or two levels removed from the perturbation were either very weak or undetectable. Top-down effects were strong when direct but attenuated quickly. Bottom-up effects were less strong but persisted as indirect effects to higher levels. Although the 'pure reciprocal' model best fits our results for the boreal forest system different models may apply to different ecosystems around the world.     

Integrating Top-Down and Bottom-Up Sensory Processing by Somato-Dendritic Interactions

The classical view of cortical information processing is that of a bottom-up process in a feedforward hierarchy. However, psychophysical, anatomical, and physiological evidence suggests that top-down effects play a crucial role in the processing of input stimuli. Not much is known about the neural mechanisms underlying these effects. Here we investigate a physiologically inspired model of two reciprocally connected cortical areas. Each area receives bottom-up as well as top-down information. This information is integrated by a mechanism that exploits recent findings on somato-dendritic interactions. (1) This results in a burst signal that is robust in the context of noise in bottom-up signals. (2) Investigating the influence of additional top-down information, priming-like effects on the processing of bottom-up input can be demonstrated. (3) In accordance with recent physiological findings, interareal coupling in low-frequency ranges is characteristically enhanced by top-down mechanisms. The proposed scheme combines a qualitative influence of top-down directed signals on the temporal dynamics of neuronal activity with a limited effect on the mean firing rate of the targeted neurons. As it gives an account of the system properties on the cellular level, it is possible to derive several experimentally testable predictions.     

The relative importance of herbivory and carnivory on the distribution of energy in a stochastic tri-trophic food web

A three-state, discrete-time Markov chain is used to model the dynamics of energy flow in a tri-trophic food web. The distribution of energy in the three trophic levels is related to the rates of flow between the trophic levels and calculated for the entire range of possible flow values. These distributions are then analysed for stability and used to test the idea that plants are resource-limited and herbivores are predation-limited. Low rates of death and decomposition, when coupled with low rates of herbivory and carnivory, tend to destabilize this food web. Food webs with higher rates of death and decomposition are relatively more stable regardless of rates of herbivory and carnivory. Plants are more prone to resource-limitation and herbivores are, in general, limited by their predators, which supports Hairston et al. (Am. Nat. 94 (1960) 421). The rate of decomposition often mediates the roles of top-down and bottom-up control of energy flow in the food web.     

Relative strengths of top-down and bottom-up forces in a tropical forest community

We tested integrative bottom-up and top-down trophic cascade hypotheses with manipulative experiments in a tropical wet forest, using the ant-plant Piper cenocladum and its associated arthropod community. We examined enhanced nutrients and light along with predator and herbivore exclusions as sources of variation in the relative biomass of plants, their herbivores (via rates of herbivory), and resident predaceous ants. The combined manipulations of secondary consumers, primary consumers, and plant resources allowed us to examine some of the direct and indirect effects on each trophic level and to determine the relative contributions of bottom-up and top-down cascades to the structure of the community. We found that enhanced plant resources (nutrients and light) had direct positive effects on plant biomass. However, we found no evidence of indirect (cascading through the herbivores) effects of plant biomass on predators or top predators. In contrast, ants had indirect effects on plant biomass by decreasing herbivory on the plants. This top-down cascade occurred whether or not plant resources were enriched, conditions which are expected to modify top-down forces. Received: 9 August 1998 / Accepted: 1 December 1998     

Is a healthy ecosystem one that is rich in parasites?

Historically, the role of parasites in ecosystem functioning has been considered trivial because a cursory examination reveals that their relative biomass is low compared with that of other trophic groups. However there is increasing evidence that parasite-mediated effects could be significant: they shape host population dynamics, alter interspecific competition, influence energy flow and appear to be important drivers of biodiversity. Indeed they influence a range of ecosystem functions and have a major effect on the structure of some food webs. Here, we consider the bottom-up and top-down processes of how parasitism influences ecosystem functioning and show that there is evidence that parasites are important for biodiversity and production; thus, we consider a healthy system to be one that is rich in parasite species.