Resorption of foliar nutrients in a regenerating southern Appalachian forest

Summary Leaves were sampled in a successional, southern Appalachian forest to estimate autumn foliar nutrient dynamics. Resorption of N and P in a successional forest equaled, or exceeded, resorption estimates for a more mature control forest. Foliar nutrient leaching was not sufficient to account for changes in autumn leaf N, P, Ca and Mg concentrations. The resorption process conserves nutrients by reducing nutrient losses from leaching and litter-fall, thereby closing the nutrient cycle in successional forests. We hypothesize that rapid recovery of primary productivity early in forest regeneration is the result of maximum nutrient resorption of limiting nutrients. Implications of these results for successional nutrient cycling theory are discussed.     

Stoichiometry of nutrient excretion by fish: interspecific variation in a hypereutrophic lake

This study investigates how nutrient cycling rates and ratios vary among fish species, with a particular focus on comparing an ecologically dominant detritivore (gizzard shad) to other fishes in a productive lake. We also examined how nutrient cycling rates are mediated by body size (as predicted by allometry theory), and how variation in nutrient cycling is related to body and food nutrient contents (according to predictions of ecological stoichiometry). As predicted by allometry, per capita nitrogen and phosphorus excretion rates increased and mass-specific excretion rates decreased, with increasing mass. Body phosphorus content was correlated with body mass only in one species, bluegill. Contrary to stoichiometric predictions, there was no relationship between body P and mass-normalized P excretion rate, or between body N:P and excreted N:P, when all individuals of all species were considered.
However, at the species level, we observed some support for a body nutrient content effect on excretion as predicted by stoichiometry theory. For example, gizzard shad had lower body P (high body N:P) and also excreted P at higher rates (lower N:P) than bluegill, which had high body P (lower body N:P). We applied the Sterner (1990) homeostatic stoichiometry model to the two most common species in the study – gizzard shad and bluegill and found that food N:P had a greater effect than consumer body N:P on excreted N:P. This indicates that, in terms of variation among these species, nutrient excretion may be more of a function of food nutrient content than the nutrient content of the consumer. These results suggest that stoichiometry can provide a framework for variation among species in nutrient cycling and for evaluating the ecosystem consequences of biodiversity loss.     

Nutrient flows between ecosystems can destabilize simple food chains

Dispersal of organisms has large effects on the dynamics and stability of populations and communities. However, current metacommunity theory largely ignores how the flows of limiting nutrients across ecosystems can influence communities. We studied a meta-ecosystem model where two autotroph-consumer communities are spatially coupled through the diffusion of the limiting nutrient. We analyzed regional and local stability, as well as spatial and temporal synchrony to elucidate the impacts of nutrient recycling and diffusion on trophic dynamics. We show that nutrient diffusion is capable of inducing asynchronous local destabilization of biotic compartments through a diffusion-induced spatiotemporal bifurcation. Nutrient recycling interacts with nutrient diffusion and influences the susceptibility of the meta-ecosystem to diffusion-induced instabilities. This interaction between nutrient recycling and transport is further shown to depend on ecosystem enrichment. It more generally emphasizes the importance of meta-ecosystem theory for predicting species persistence and distribution in managed ecosystems.     

Feedbacks of consumer nutrient recycling on producer biomass and stoichiometry: separating direct and indirect effects

Herbivores can have both direct (consumptive) and indirect (nutrient‐mediated) effects on primary producer biomass and nutrient stoichiometry. Ecological stoichiometry theory predicts that herbivores of contrasting body stoichiometry will differentially remineralize nutrients, resulting in feedbacks on producer stoichiometry. We experimentally separated direct and indirect effects of aquatic vertebrate grazers on periphyton by manipulating grazer abundance and identity in mesocosms, and using grazer exclusion cages to expose periphyton to recycled nutrients in the absence of direct grazing. In experiment 1, we used a catfish with high body phosphorus (low body N:P), Ancistrus triradiatus, to assess consumptive versus nutrient‐mediated effects of grazer density on periphyton. In experiment 2, we compared the nutrient‐mediated effects of grazing by Ancistrus triradiatus and Rana palmipes, a tadpole with low body phosphorus and high body N:P. In experiment 1, we found that increasing catfish density led to lower biomass and particulate nutrients in periphyton through direct consumptive effects, but that nutrient‐mediated indirect effects enhanced periphyton biomass when grazers were experimentally separated from direct contact with periphyton. As predicted by stoichiometry theory, nutrient recycling by this P‐rich grazer tended to increase algal C:P and N:P (although effects were not statistically significant), while their consumptive effects reduced algal C:P and N:P. In experiment 2, grazer identity had strong effects on dissolved water nutrient concentrations, N recycling (measured with a ¹⁵N tracer), and periphyton stoichiometry. In accordance with stoichiometry theory, catfish increased N concentrations and recycling rates leading to higher periphyton N:P, while tadpoles had greater effects on P availability leading to lower periphyton N:P. Our experiments elucidate the importance of both the density and identity of grazers in controlling periphyton biomass and stoichiometry through consumptive and nutrient‐mediated effects, and support the power of ecological stoichiometry theory to predict feedbacks on producer stroichiometry arising from consumer stoichiometry through nutrient recycling.     

Stability and stabilization for models of chemostats with multiple limiting substrates

We study chemostat models in which multiple species compete for two or more limiting nutrients. First, we consider the case where the nutrient flow and species removal rates and input nutrient concentrations are all given as positive constants. In that case, we use Brouwer degree theory to give conditions guaranteeing that the models admit globally asymptotically stable componentwise positive equilibrium points, from all componentwise positive initial states. Then we use the results to develop stabilization theory for a class of controlled chemostats with two or more limiting nutrients. For cases where the dilution rate and input nutrient concentrations can be selected as controls, we prove that many different componentwise positive equilibria can be made globally asymptotically stable. This extends the existing control results for chemostats with one limiting nutrient. We demonstrate our methods in simulations.     

Stability and stabilization for models of chemostats with multiple limiting substrates

We study chemostat models in which multiple species compete for two or more limiting nutrients. First, we consider the case where the nutrient flow and species removal rates and input nutrient concentrations are all given as positive constants. In that case, we use Brouwer degree theory to give conditions guaranteeing that the models admit globally asymptotically stable componentwise positive equilibrium points, from all componentwise positive initial states. Then we use the results to develop stabilization theory for a class of controlled chemostats with two or more limiting nutrients. For cases where the dilution rate and input nutrient concentrations can be selected as controls, we prove that many different componentwise positive equilibria can be made globally asymptotically stable. This extends the existing control results for chemostats with one limiting nutrient. We demonstrate our methods in simulations.     

Quasi-equilibrium reduction in a general class of stoichiometric producer-consumer models

This article compares a general closed nutrient, stoichiometric producer-consumer model to a two-dimensional 'quasi-equilibrium' approximation. We demonstrate that the quasi-equilibrium system can be rigorously analysed, resulting in nullcline-based criteria for the local stability of system equilibria and for the non-existence of periodic orbits. These results are applied to a study of the dependence of the reduced system on nutrient and energy enrichment. When energy and nutrient enrichment are considered together, the associated bifurcation structures of the two models are seen to share the same essential qualitative characteristics. However, numerical simulations of the three-dimensional parent model show highly complex domains of the persistence and extinction that by Poincare-Bendixson theory are not possible for the two-dimensional reduction. This complexity demonstrates a major difference between the two models, and suggests potential challenges in the use of either model for predicting the long-term behaviour of real-world systems at specific nutrient and energy levels.     

Quasi-equilibrium reduction in a general class of stoichiometric producer–consumer models

This article compares a general closed nutrient, stoichiometric producer–consumer model to a two-dimensional ‘quasi-equilibrium’ approximation. We demonstrate that the quasi-equilibrium system can be rigorously analysed, resulting in nullcline-based criteria for the local stability of system equilibria and for the non-existence of periodic orbits. These results are applied to a study of the dependence of the reduced system on nutrient and energy enrichment. When energy and nutrient enrichment are considered together, the associated bifurcation structures of the two models are seen to share the same essential qualitative characteristics. However, numerical simulations of the three-dimensional parent model show highly complex domains of the persistence and extinction that by Poincare–Bendixson theory are not possible for the two-dimensional reduction. This complexity demonstrates a major difference between the two models, and suggests potential challenges in the use of either model for predicting the long-term behaviour of real-world systems at specific nutrient and energy levels.     

Theories and methods on plant nutrition and growth

A theory comprising two basic concepts relating nutrition and growth is presented. The first concept is a nutrient flux model and is based upon studies of plants at constant internal nutrient concentrations, where a formal mathematical derivation shows that the relative uptake rate (R_U) and the relative growth rate (R_G) are equal. Deviations from equality are results of experimental insufficiencies and errors. The second concept is based on the observation that R_G is linearly related to the internal nutrient concentration. The slope represents nutrient productivity (P_n), an important parameter expressing growth rate per unit of nutrient. Light and the plant genome, for example, influence the value of the proportionality factor, P_n, but not the formal relationship between the internal nutrient concentration and R_G Not only the theory itself but many results and conclusions are very different from those obtained with traditional methods. In experiments where R_U was controlled during the exponential period of growth, the relationships between treatment (the relative addition rate, R_A), nutrient uptake (R_U) and growth (R_G) were reproduced with extremely low variability. In agreement with theory, internal nutrient concentration and R_G remained stable over time (steady-state). An extension of the theory is based upon the empirical assumption that after exponential growth, self-shading and ageing reduce P_n in proportion to biomass. This assumption has been successfully applied in predicting growth of forest stands, but the nature of the growth reduction is little understood. The generalized model has few parameters and can easily be rewritten to suit different experimental aims, for example to establish reference values and to model changes in soil fertility. Further extension and understanding of the model and different interpretations of the parameters are discussed.     

The nutrient-load hypothesis: patterns of resource limitation and community structure driven by competition for nutrients and light

Resource competition theory predicts that the outcome of competition for two nutrients depends on the ratio at which these nutrients are supplied. Yet there is considerable debate whether nutrient ratios or absolute nutrient loads determine the species composition of phytoplankton and plant communities. Here we extend the classical resource competition model for two nutrients by including light as additional resource. Our results suggest the nutrient-load hypothesis, which predicts that nutrient ratios determine the species composition in oligotrophic environments, whereas nutrient loads are decisive in eutrophic environments. The underlying mechanism is that nutrient enrichment shifts the species interactions from competition for nutrients to competition for light, which favors the dominance of superior light competitors overshadowing all other species. Intermediate nutrient loads can generate high biodiversity through a fine-grained patchwork of two-species and three-species coexistence equilibria. Depending on the species traits, however, competition for nutrients and light may also produce multiple alternative stable states, suppressing the predictability of the species composition. The nutrient-load hypothesis offers a solution for several discrepancies between classical resource competition theory and field observations, explains why eutrophication often leads to diversity loss, and provides a simple conceptual framework for patterns of biodiversity and community structure observed in nature.     

Roots in space: a spatially explicit model for below-ground competition in plants

Game theory provides an untapped framework for predicting how below-ground competition will influence root proliferation in a spatially explicit environment. We model root competition for space as an evolutionary game. In response to nutrient competition between plants, an individual's optimal strategy (the spatial distribution of root proliferation) depends on the rooting strategies of neighbouring plants. The model defines and predicts the fundamental (in the absence of competition) and realized (in the presence of competition) root space of an individual plant. Overlapping fundamental root spaces guarantee smaller, yet still overlapping, realized root spaces as individuals concede some but not all space to a neighbour's roots. Root overlap becomes an intentional consequence of the neighbouring plants playing a nutrient foraging game. Root proliferation and regions of root overlap should increase with soil fertility, decline with the distance cost of root production (e.g. soil compactness) and shift with competitive asymmetries. Seemingly erratic patterns of root proliferation and root overlap become the expected outcome of nutrient foraging games played in soils with small-scale heterogeneities in nutrient availability.     

The ontogenetic stoichiometric bottleneck stabilizes herbivore–autotroph dynamics

In the ecological stoichiometry theory of population dynamics, ontogenetic changes in nutrient demand have been ignored. Here, I studied a stage-structured Daphnia–algae herbivore–autotroph model, in which the juveniles of the herbivore had a higher nutrient (phosphorous) demand for maturation than the adults for reproduction. The model predicted that while an increase in the juvenile nutrient demand (i.e., ontogenetic stoichiometric bottleneck) affects stage-specific performances in complex ways through nutrient dynamics and resource quality, in general it has stabilizing effects on the population dynamics.     

Animal pee in the sea: consumer‐mediated nutrient dynamics in the world's changing oceans

Humans have drastically altered the abundance of animals in marine ecosystems via exploitation. Reduced abundance can destabilize food webs, leading to cascading indirect effects that dramatically reorganize community structure and shift ecosystem function. However, the additional implications of these top‐down changes for biogeochemical cycles via consumer‐mediated nutrient dynamics (CND) are often overlooked in marine systems, particularly in coastal areas. Here, we review research that underscores the importance of this bottom‐up control at local, regional, and global scales in coastal marine ecosystems, and the potential implications of anthropogenic change to fundamentally alter these processes. We focus attention on the two primary ways consumers affect nutrient dynamics, with emphasis on implications for the nutrient capacity of ecosystems: (1) the storage and retention of nutrients in biomass, and (2) the supply of nutrients via excretion and egestion. Nutrient storage in consumer biomass may be especially important in many marine ecosystems because consumers, as opposed to producers, often dominate organismal biomass. As for nutrient supply, we emphasize how consumers enhance primary production through both press and pulse dynamics. Looking forward, we explore the importance of CDN for improving theory (e.g., ecological stoichiometry, metabolic theory, and biodiversity–ecosystem function relationships), all in the context of global environmental change. Increasing research focus on CND will likely transform our perspectives on how consumers affect the functioning of marine ecosystems.     

“生态平衡施肥”的理论基础和技术体系

施肥是与产量、品质、成本、土壤培肥、污染等问题密切相关的复合生态系统物质循环调控的重要措施,以实现经济效益、生态效益和社会效益的综合效益最佳为目标,建立了“生态平衡施肥”的理论基础和技术体系,结合国情提出“生态平衡施肥’实现的关键条件和具体模式。“生态平衡施肥”是以施肥系统养分循环原理－－“通用施肥模型”为理论基础,以衫和和高新技术化组装的技术体系为实现手段,以“生态型肥料”为载体的新的施肥系统。     

Optimal foraging for multiple nutrients in an unpredictable environment

Foraging theory has typically been concerned with the acquisition of a single resource even though organisms from mammals to protozoa are capable of balancing their requirements for multiple resources. Existing theory concerning multiple nutrients from multiple foods does not predict the sequence of food selection. We constructed an optimisation model of the simplest case of two foods containing differing amounts of two nutrients. We begin with the well-supported assumption that reproductive value declines with the distance from target nutrient intake. We show that nutrient space divides into two distinct areas where the animal should exclusively consume one food or the other. The organism thus initially concentrates on one food type until the border between the areas is reached and then moves as closely as possible along the border to approach the target. This strategy is commonly observed in a range of organisms, suggesting that the assumed fitness function is common.     

Do nutrient-competition models predict nutrient availabilities in limnetic ecosystems?

Recent theory on resource competition, predicated on the importance of hypothesized trade-offs between minimum requirements for nutrient resources, predicts that there should be negative correlations between the supply rate of major limiting nutrients and the availability of at least some secondary nutrients and/or among the availabilities of different limiting nutrients. However, an analysis of four data sets from large-scale surveys of lakes shows mostly positive correlations among the availabilities and supplies of nutrients. In contrast, a fifth data set, obtained in an area of high acidification, does show several important negative correlations that are consistent with the nutrient competition models. Further analyses suggest two possible explanations for the preponderance of positive correlation. Negative correlations between nutrients and light indicate that an important trade-off among species regulating phytoplankton may involve low light requirements versus low nutrient requirements. The existence of negative correlations in nutrient availabilities in acidic lakes (where herbivory appears less important than in buffered lakes) also suggests that another important trade-off may involve an ability to minimize loss rates (especially due to grazing) versus an overall ability to exploit nutrient resources. Received: 23 February 1996 / Accepted: 30 September 1996     

可持续的污水处理过程与展望

传统上,污水在处理过程中其所含有的能源和营养物质等也被消耗了。可持续的水处理不仅包括污水无害化处理,还包括能源和营养物的综合回收利用。综合论述了可持续的污水处理概念、理论、方法和污水处理过程中物质回收技术的发展。     

The effect of heterogeneity on invasion in spatial epidemics: from theory to experimental evidence in a model system

Heterogeneity in host populations is an important factor affecting the ability of a pathogen to invade, yet the quantitative investigation of its effects on epidemic spread is still an open problem. In this paper, we test recent theoretical results, which extend the established "percolation paradigm" to the spread of a pathogen in discrete heterogeneous host populations. In particular, we test the hypothesis that the probability of epidemic invasion decreases when host heterogeneity is increased. We use replicated experimental microcosms, in which the ubiquitous pathogenic fungus Rhizoctonia solani grows through a population of discrete nutrient sites on a lattice, with nutrient sites representing hosts. The degree of host heterogeneity within different populations is adjusted by changing the proportion and the nutrient concentration of nutrient sites. The experimental data are analysed via Bayesian inference methods, estimating pathogen transmission parameters for each individual population. We find a significant, negative correlation between heterogeneity and the probability of pathogen invasion, thereby validating the theory. The value of the correlation is also in remarkably good agreement with the theoretical predictions. We briefly discuss how our results can be exploited in the design and implementation of disease control strategies.     

Growth rates of yeast colonies on solid media

Previous measurements of growth rates of giant yeast colonies on solid media are shown to be unreliable as they depend strongly on extraneous factors such as the proximity of other colonies and the dimensions of the apparatus used. The hitherto unexplained dependence of the growth rate on the square root of the growth limiting nutrient concentration is explained by constructing a theory based on the diffusion of nutrient towards the colony which makes use of many ideas used in the theory of flame propagation. The theory also explains why the temperature dependence of the homogeneous growth constant is different from that observed in the surface colony, and it requires the existence of a lag phase in the homogeneous culture kinetics if the velocity of propagation of the culture is to be independent of inoculum size and shape. Both phenomena are known to occur.     

Detritivory and the stoichiometry of nutrient cycling by a dominant fish species in lakes of varying productivity

Little is known about the stoichiometry of nutrient cycling by detritivores. Therefore, we explored stoichiometric relationships in an omnivorous/detritivorous fish (gizzard shad, Dorosoma cepedianum) in three lakes that differed in productivity. Gizzard shad can feed on plankton and sediment detritus, but in all three lakes adult gizzard shad derived >98% of carbon (C) and phosphorus (P), and >90% of nitrogen (N) from sediment detritus, and the remainder from zooplankton.
Gizzard shad selectively consumed detritus with higher C, N and P concentrations than ambient lake sediments. Selective detritivory (i.e. the nutrient content of consumed detritus divided by the nutrient content of ambient detritus) was most pronounced in the lake with the lowest detrital nutrient concentrations. N and P cycling rates per fish were also consistently higher in this lake, in agreement with the prediction of stoichiometry theory that excretion rates should increase with food nutrient content. Among-lake differences in nutrient cycling rates were unrelated to inter-lake variation in fish body nutrient contents, which was minimal. The N:P ratio excreted was near Redfield (∼14:1) in all three lakes.
Stoichiometric analyses showed that the C:N and C:P ratios of sediment detritus were much higher (∼2.8×) than ratios of gizzard shad bodies, revealing substantial N and P imbalances between consumers and their food source. Gizzard shad alleviate N imbalance by selectively feeding on high N detritus (low C:N, high N:P), and apparently alleviate P imbalance by excreting nutrients at a higher N:P than that of their food or their bodies. Thus, this detritivore apparently regulates nutrient acquisition and allocation via both pre-absorption processes (selective feeding) and post-absorptive processes (differential N and P excretion).