富食悖论研究进展

Michael Rosenzweig于1971年首次提出＂富食悖论＂这一概念：在简单的被捕食者-捕食者系统中,随着营养物质供应的增加,系统变得不再稳定,并产生大振幅波动,最终导致系统内的物种灭亡。然而,许多实验结果并不支持Rosenzweig的理论。综述＂富食悖论＂的定义、理论和实验研究,同时综述各种解释实际与理论之间矛盾的机制,最后探讨＂富食悖论＂的研究前景。     

The paradox of enrichment in an adaptive world

Paradoxically, enrichment can destabilize a predator-prey food web. While adaptive dynamics can greatly influence the stability of interaction systems, few theoretical studies have examined the effect of the adaptive dynamics of interaction-related traits on the possibility of resolution of the paradox of enrichment. We consider the evolution of attack and defence traits of a predator and two prey species in a one predator-two prey system in which the predator practises optimal diet use. The results showed that optimal foraging alone cannot eliminate a pattern of destabilization with enrichment, but trait evolution of the predator or prey can change the pattern to one of stabilization, implying a possible resolution of the paradox of enrichment. Furthermore, trait evolution in all species can broaden the parameter range of stabilization. Importantly, rapid evolution can stabilize this system, but weaken its stability in the face of enrichment.     

Interplay between the paradox of enrichment and nutrient cycling in food webs

Nutrient cycling is fundamental to ecosystem functioning. Despite recent major advances in the understanding of complex food web dynamics, food web models have so far generally ignored nutrient cycling. However, nutrient cycling is expected to strongly impact food web stability and functioning. To make up for this gap, we built an allometric and size structured food web model including nutrient cycling. By releasing mineral nutrients, recycling increases the availability of limiting resources for primary producers and links each trophic level to the bottom of food webs. We found that nutrient cycling can provide a significant part of the total nutrient supply of the food web, leading to a strong enrichment effect that promotes species persistence in nutrient poor ecosystems but leads to a paradox of enrichment at high nutrient inputs. The presence of recycling loops linking each trophic level to the basal resources weakly affects species biomass temporal variability in the food web. Recycling loops tend to slightly dampen the destabilising effect of nutrient enrichment on consumer temporal variability while they have opposite effects for primary producers. By considering nutrient cycling, this new model improves our understanding of the response of food webs to nutrient availability and opens perspectives to better link studies on food web dynamics and ecosystem functioning.     

Dispersal delays, predator-prey stability, and the paradox of enrichment

It takes time for individuals to move from place to place. This travel time can be incorporated into metapopulation models via a delay in the interpatch migration term. Such a term has been shown to stabilize the positive equilibrium of the classical Lotka-Volterra predator-prey system with one species (either the predator or the prey) dispersing. We study a more realistic, Rosenzweig-MacArthur, model that includes a carrying capacity for the prey, and saturating functional response for the predator. We show that dispersal delays can stabilize the predator-prey equilibrium point despite the presence of a Type II functional response that is known to be destabilizing. We also show that dispersal delays reduce the amplitude of oscillations when the equilibrium is unstable, and therefore may help resolve the paradox of enrichment.     

Towards resolving the paradox of enrichment: the impact of zooplankton vertical migrations on plankton systems stability

Eutrophication, often resulting from human activity, is a serious threat to aquatic communities. Theoretical analysis of this phenomenon, based on conceptual mathematical models, leads to controversial predictions known as Rosenzweig's paradox of enrichment. At the same time, field observations demonstrate that real plankton communities exhibit various mechanisms of self-regulation which can buffer negative effects of enrichment. In this paper, we study potential effects of zooplankton vertical migration on stability of plankton systems functioning. We consider an intrinsically unstable plankton model, which is characterized by an unlimited phytoplankton multiplication and population oscillations of increasing amplitude, and investigate whether vertical migrations of zooplankton can stabilize such a system at low plankton densities. By means of developing two different models accounting for different ecological situations, e.g. deep waters and shallow waters, we show that vertical migrations of zooplankton can result in stabilization of eutrophic plankton systems. Thus, we show that this mechanism, rarely taken into account in models of plankton dynamics, may be important for resolving the paradox of enrichment in plankton communities.     

Discrete consumers,small scale resource heterogeneity,and population stability

We present a consumer-resource model in which individual consumers subsist on a continuum of resource distributed over a very large number of small “bite-sized” patches, each patch being sufficiently small that all its resource is eaten whenever a consumer visits. This form of consumer–resource interaction forces a heterogeneous distribution of resource among the patches, and may dampen out the large amplitude, consumer-resource cycles that are predicted by traditional models of well-mixed, spatially homogeneous systems. The resource equilibrium does not increase with enrichment, a prediction that distinguishes this model from models that invoke direct or indirect consumer density dependence as a stabilizing mechanism.     

Imperfect optimal foraging and the paradox of enrichment

We show that the paradox of enrichment can be theoretically resolved in a flexible predator–prey system in which the predator practices imperfect optimal foraging. A previous study showed that perfect optimal foraging can mitigate increases in the amplitude of population oscillations associated with enrichment, but it did not show a stabilization pattern. Our results show that imperfect optimal foraging can stabilize the system and resolve the paradox of enrichment under nonequilibrium dynamics. Furthermore, the degree of stabilization with enrichment was stronger when the imperfection of optimal foraging was larger.     

Response to enrichment, type and timing: small mammals vary in their response to a springtime cicada but not a carbohydrate pulse

1. Masting events in the autumn provide a carbohydrate-rich pulse of resources that can influence the dynamics of small mammals and their natural enemies. Similar patterns are observed with the periodical cicada emergence which provides a protein-rich pulse in the spring, but comparisons are confounded by timing and food type. 2. We compared the influence of a naturally occurring spring pulse of cicadas with an experimental spring pulse of carbohydrate-rich seeds. We used a replicated population level field experiment and capture-mark-recapture techniques to record the vital rates, demographics, and abundance of Peromyscus leucopus (the white-footed mouse), as well as other small mammals and their parasites. 3. The density of P. leucopus on grids where cicadas emerged was 55% higher than controls as a consequence of early breeding. This was followed by an increase in the prevalence of the nematode Pterygodermatities peromysci, reduced breeding and decreased recruitment rates. Other small mammals including Tamias striatus (eastern chipmunk) and Blarina brevicauda (short-tailed shrew), increased in density, but there was no affect on Sorex cinereus (masked shrew). 4. In contrast to the presence of cicadas, there was no influence of sunflower seed supplementation on small mammal density, vital rates, or reproduction with the exception of an increase in B. brevicauda density. The response of small mammals to seasonal pulses depends on timing, food type, and species.     

Primate puzzleboard: A simple environmental enrichment device for captive chimpanzees

An environmental enrichment device was designed and evaluated for captive chimpanzees. The puzzleboard consisted of a 61-cm, square piece of clear plastic with holes of various sizes drilled into the center. The puzzleboard was bolted to the roof of a chimpanzee enclosure. Flavored primate treats were arranged on top of the board, and the chimpanzee used a finger to manipulate the food across the board to be retrieved through the larger holes. The behavior and puzzle use of 29 chimpanzees housed in eight groups were evaluated while the device was empty and while filled with food. For each group, four 1-hr scan samples were recorded for each condition. Data showed that an average of 40 min elapsed from the time the puzzleboard was filled to the last recorded contact by a subject. Mean individual use ranged from 4.8 to 23.8 min/hr. Female chimpanzees used the puzzle significantly more often than did males. Aggressive, affiliative, inactive, and self-directed behaviors were significantly reduced when the puzzle was filled. In addition, the puzzleboard was inexpensive, sturdy, and easily constructed, mounted, and operated.     

The dynamics of food chains under climate change and nutrient enrichment

Warming has profound effects on biological rates such as metabolism, growth, feeding and death of organisms, eventually affecting their ability to survive. Using a nonlinear bioenergetic population-dynamic model that accounts for temperature and body-mass dependencies of biological rates, we analysed the individual and interactive effects of increasing temperature and nutrient enrichment on the dynamics of a three-species food chain. At low temperatures, warming counteracts the destabilizing effects of enrichment by both bottom-up (via the carrying capacity) and top-down (via biological rates) mechanisms. Together with increasing consumer body masses, warming increases the system tolerance to fertilization. Simultaneously, warming increases the risk of starvation for large species in low-fertility systems. This effect can be counteracted by increased fertilization. In combination, therefore, two main drivers of global change and biodiversity loss can have positive and negative effects on food chain stability. Our model incorporates the most recent empirical data and may thus be used as the basis for more complex forecasting models incorporating food-web structure.     

Biomass diversity and stability of food webs in aquatic ecosystems

The diversity–stability hypothesis in ecology asserts that biodiversity begets stability of ecological systems. This hypothesis has been supported by field studies on primary producers in grasslands, in which the interaction between species is mostly competition. As to ecosystems with multitrophic predatory interaction, however, no definite consensus has been arrived at for the relation between trophic diversity and ecosystem stability. The stability index suitable to ecosystems with predatory interaction is given by MacArthurs idea of stability and its formulation by Rutledge et al. More suitable indices of stability (relative conditional entropy) are proposed in this study for the comparison of different ecosystems, and the validity of the diversity–stability hypothesis for food webs (networks of predation) with many trophic compartments in natural aquatic ecosystems is examined. Results reveal that an increase in the biomass diversity of trophic compartments causes an increase in the whole systemic stability of food webs in aquatic ecosystems. Hence, evidence of the whole systemic validity of the diversity–stability hypothesis for natural aquatic ecosystems with ubiquitous multitrophic predatory interaction was obtained for the first time.     

A brief note on the effects of novel enrichment on an unwanted behaviour of captive bonobos

Animals in captivity may show undesirable behaviour when they are not sufficiently challenged, making it essential for their well-being to provide daily enrichment to zoo animals. Primates need a regular replacement with novel enrichment objects to prevent them from getting bored and look for challenging situations which might be even more undesirable. A socially housed group of bonobos showed behavioural problems such as stealing and rough handling and playing with a newborn infant in the group. To study whether providing novel environmental enrichment has an effect on this phenomenon we performed an experiment where we observed the change in behaviour in the presence of familiar and novel enrichment devices. When provided with novel enrichments, the general activity of the group increased. Simultaneously, these novel behavioural challenges significantly decreased the frequency of taking the newborn infant from its mother. Our results confirm previous findings that continuous application of new enrichments is necessary for providing sensory stimulation to primates as it promotes their well-being and indirectly might influence breeding success as well.     

A resolution of the paradox of enrichment

Theoretical studies have shown a paradoxical destabilizing response of predator-prey ecosystems to enrichment, but there is the gap between the intuitive view of nature and this theoretical prediction. We studied a minimal predator-prey system (a two predator-two prey system) in which the paradox of enrichment pattern can vanish; the destabilization with enrichment is reversed, leading to stabilization (a decrease in the amplitude of oscillation of population densities). For resolution of the paradox, two conditions must be met: (1) the same prey species must be preferred as a dietary item by both predator species, creating the potential for high exploitative competition between the predator species, and (2), while both predators are assumed to select their diet in accordance with optimal diet utilization theory, one predator must be a specialist and the other a generalist. In this system, the presence of a less profitable prey species can cause the increase in population oscillation amplitudes associated with increasing enrichment to be suppressed via the optimal diet utilization of the generalist predator. The resulting stabilization is explained by the mitigating effect of the less profitable prey showing better population growth with increasing enrichment on the destabilization underlying the specialist predator and prey relation, thus resolving the paradox of enrichment.     

Environmental variability uncovers disruptive effects of species' interactions on population dynamics

How species respond to changes in environmental variability has been shown for single species, but the question remains whether these results are transferable to species when incorporated in ecological communities. Here, we address this issue by analysing the same species exposed to a range of environmental variabilities when (i) isolated or (ii) embedded in a food web. We find that all species in food webs exposed to temporally uncorrelated environments (white noise) show the same type of dynamics as isolated species, whereas species in food webs exposed to positively autocorrelated environments (red noise) can respond completely differently compared with isolated species. This is owing to species following their equilibrium densities in a positively autocorrelated environment that in turn enables species–species interactions to come into play. Our results give new insights into species'' response to environmental variation. They especially highlight the importance of considering both species'' interactions and environmental autocorrelation when studying population dynamics in a fluctuating environment.     

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.     

A solution to the human paradox: fundamental ontogenies and heterochronies

Solving the human paradox means explaining how a genetic difference of a mere 1% can be consistent with 5 million years of anatomical transformation from great apes to present-dayHomo sapiens. The solution proposed here is that of the internal history of ontogenetic change. A concept of “fundamental ontogeny” is developed and deduced from comparison between living and fossil primates. The fossil human lineage can be summarized into five fundamental ontogenies corresponding to successive skull plans (bauplans) resulting from five major phases of craniofacial contraction: prosimians (adapiforms), monkey apes (propliopithecidae), great apes (dryopithecidae), australopithecines andHomo. The morphological areas defined by these skull plans include more-or-less numerous species. This concept leads to renewed debate about (i) the relationship between speciation and bauplans, and (ii) the mechanisms involved in the successive steps of cranio-facial contraction and the correlated morphological changes. It is suggested that, from great apes to modern man, numerous heterochronies (hypermorphosis, hypomorphosis and post-displacements) have occurred during ontogeny, allowing the acquisition of permanent bipedalism inAustralopithecus andHomo, the increased cranial capacity of primitive forms ofHomo, and the disappearance of simian characters associated with renewed increase in cranial capacity inH. sapiens.     

The influence of nutrient enrichment on riverine food web function and stability

Nutrient enrichment of rivers and lakes has been increasing rapidly over the past few decades, primarily because of agricultural intensification. Although nutrient enrichment is known to drive excessive algal and microbial growth, which can directly and indirectly change the ecological community composition, the resulting changes in food web emergent properties are poorly understood. We used ecological network analysis (ENA) to examine the emergent properties of 12 riverine food webs across a nutrient enrichment gradient in the Manawatu, New Zealand. We also derive Keystone Sensitivity Indices to explore whether nutrients change the trophic importance of species in a way that alters the resilience of the communities to further nutrient enrichment or floods. Nutrient enrichment resulted in communities composed of energy inefficient species with high community (excluding microbes) respiration. Community respiration was several times greater in enriched communities, and this may drive hypoxic conditions even without concomitant changes in microbial respiration. Enriched communities exhibited weaker trophic cascades, which may yield greater robustness to energy flow loss. Interestingly, enriched communities were also more structurally and functionally affected by species sensitive to flow disturbance making these communities more vulnerable to floods.     

The protozoon and the cell: A brief twentieth-century overview

This paper represents the subject matter of an invitational presentation made October 24, 1986, at the meetings of the History of Science Society held in Pittsburgh, Pa. By uncanny coincidence, considering the topics of our papers, that day was also the birthday (the 354th) of the founding father of protozoology, Antony van Leeuwenhoek! Appreciation is expressed here for the support of National Science Foundation grant BSR 83-07113.