Quantity–quality trade‐offs revealed using a multiscale test of herbivore resource selection on elemental landscapes

1. Herbivores consider the variation of forage qualities (nutritional content and digestibility) as well as quantities (biomass) when foraging. Such selection patterns may change based on the scale of foraging, particularly in the case of ungulates that forage at many scales.
2. To test selection for quality and quantity in free‐ranging herbivores across scales, however, we must first develop landscape‐wide quantitative estimates of both forage quantity and quality. Stoichiometric distribution models (StDMs) bring opportunity to address this because they predict the elemental measures and stoichiometry of resources at landscape extents.
3. Here, we use StDMs to predict elemental measures of understory white birch quality (% nitrogen) and quantity (g carbon/m²) across two boreal landscapes. We analyzed global positioning system (GPS) collared moose (n = 14) selection for forage quantity and quality at the landscape, home range, and patch extents using both individual and pooled resource selection analyses. We predicted that as the scale of resource selection decreased from the landscape to the patch, selection for white birch quantity would decrease and selection for quality would increase.
4. Counter to our prediction, pooled‐models showed selection for our estimates of quantity and quality to be neutral with low explanatory power and no scalar trends. At the individual‐level, however, we found evidence for quality and quantity trade‐offs, most notably at the home‐range scale where resource selection models explain the largest amount of variation in selection. Furthermore, individuals did not follow the same trade‐off tactic, with some preferring forage quantity over quality and vice versa.
5. Such individual trade‐offs show that moose may be flexible in attaining a limiting nutrient. Our findings suggest that herbivores may respond to forage elemental compositions and quantities, giving tools like StDMs merit toward animal ecology applications. The integration of StDMs and animal movement data represents a promising avenue for progress in the field of zoogeochemistry.

     

Spatial stoichiometry: cross‐ecosystem material flows and their impact on recipient ecosystems and organisms

Cross‐ecosystem material flows, in the form of inorganic nutrients, detritus and organisms, spatially connect ecosystems and impact food web dynamics. To date research on material flows has focused on the impact of the quantity of these flows and largely ignored their elemental composition, or quality. However, the ratios of elements like carbon, nitrogen and phosphorus can influence the impact material flows have on food web interactions through stoichiometric mismatches between resources and consumers. The type and movement of materials likely vary in their ability to change stoichiometric constraints within the recipient ecosystem and materials may undergo changes in their own stoichiometry during transport. In this literature review we evaluate the importance of cross‐ecosystem material flows within the framework of ecological stoichiometry. We explore how movement in space and time impacts the stoichiometry of material flow, as these transformations are essential to consider when assessing the ability of these flows to impact food web productivity and ecosystem functioning. Our review suggests that stoichiometry of cross‐ecosystem material flows are highly dynamic and undergo changes during transport across the landscape or from human influence. These material flows can impact recipient organisms if they change stoichiometry of the abiotic medium, or provide resources that have a different stoichiometry to in situ resources. They might also alter consumer excretion rates, in turn altering the availability of nutrients in the recipient ecosystem. These alterations in stoichiometric constraints of recipient organisms can have cascading trophic effects and shape food web dynamics. We highlight significant gaps in the literature and suggest new avenues for research that explore how cross‐ecosystem material flows impact recipient ecosystems when considering differences in stoichiometric quality, their movement through the landscape and across ecosystem boundaries, and the nutritional constraints of the recipient organisms.     

Woodstoich III: Integrating tools of nutritional geometry and ecological stoichiometry to advance nutrient budgeting and the prediction of consumer‐driven nutrient recycling

Within the last two decades, ecological stoichiometry (ES) and nutritional geometry (NG, also known as geometric framework for nutrition) have delivered novel insights into core questions of nutritional ecology. These two nutritionally explicit frameworks differ in the ‘nutrient currency’ used and the focus of their past research; behavioural feeding strategies in NG, mainly investigating terrestrial organisms, and trophic ecology in ES, mainly in aquatic settings. However, both NG and ES have developed in explaining patterns across various scales of biological organization. Integrating specific tools of these frameworks could advance the field of nutritional ecology by unifying theoretical and empirical approaches from the organismal to ecosystem level processes. Toward this integration, we identified 1) nutrient/element budgets as a shared concept of both frameworks that encompass nutrient intake, retention, and release, 2) response surface plots of NG as powerful tools to illustrate processes at the organismal level and 3) the concept of consumer‐driven nutrient recycling (CNR) of ES as a useful tool bridging organism and ecosystem scales. We applied response surface plots to element budget data from an ES study to show how this approach can deliver new insights at the organismal level, e.g. by showing the interplay between egestion and excretion depending simultaneously on the consumed amount of carbon and phosphorus based on variation across individuals. By integrating concepts of ES and NG to model microbial uptake and mineralization of nitrogenous wastes reported in a NG study, we also demonstrate that considering biochemically explicit mineralization rates of organic wastes can improve predictions of CNR by reducing over‐ or underestimation of mineralization depending on the quality of the consumer's diet. Our presented tools and approaches can help to bridge the organismal and ecosystem level, advancing the predictive power of studies in nutritional ecology at multiple ecological scales.     

Divergence in plant and microbial allocation strategies explains continental patterns in microbial allocation and biogeochemical fluxes

Allocation trade‐offs shape ecological and biogeochemical phenomena at local to global scale. Plant allocation strategies drive major changes in ecosystem carbon cycling. Microbial allocation to enzymes that decompose carbon vs. organic nutrients may similarly affect ecosystem carbon cycling. Current solutions to this allocation problem prioritise stoichiometric tradeoffs implemented in plant ecology. These solutions may not maximise microbial growth and fitness under all conditions, because organic nutrients are also a significant carbon resource for microbes. I created multiple allocation frameworks and simulated microbial growth using a microbial explicit biogeochemical model. I demonstrate that prioritising stoichiometric trade‐offs does not optimise microbial allocation, while exploiting organic nutrients as carbon resources does. Analysis of continental‐scale enzyme data supports the allocation patterns predicted by this framework, and modelling suggests large deviations in soil C loss based on which strategy is implemented. Therefore, understanding microbial allocation strategies will likely improve our understanding of carbon cycling and climate.     

A conceptual framework for ecosystem stoichiometry: balancing resource supply and demand

The development of ecological stoichiometry has centered on organisms and their interactions, with less emphasis on the meaning or value of a comprehensive ecosystem stoichiometry at larger scales. Here we develop a conceptual framework that relates internal processes and exogenous factors in spatially- and temporally-linked ecosystems. This framework emerges from a functional view of ecosystem stoichiometry rooted in understanding the causes and consequences of relative stoichiometric balance, defined as the balance between ratios of resource supply and demand. We begin by modifying a graphical model based on resource ratio competition theory that relates resource supply and demand to ecosystem processes. This approach identified mechanisms, or stoichiometric schemes, through which ecosystems respond to variable resource supply. We expand this view by considering the effects of exogenous factors other then resource supply that comprise a stoichiometric template that influences stoichiometric balance within ecosystems. We then describe a number of examples of patterns in organismal stoichiometry in several types of ecosystems that illustrate stoichiometric schemes and factors that impinge directly on stoichiometric patterns. Next, we conduct an initial analysis of the stoichiometric effects of spatial linkages between ecosystems, and how those relate to boundary dynamics and hot spot development. We conclude by outlining research directions that will significantly advance our understanding of stoichiometric constraints on ecosystem structure and function.     

Environmental and organismal predictors of intraspecific variation in the stoichiometry of a neotropical freshwater fish

The elemental composition of animals, or their organismal stoichiometry, is thought to constrain their contribution to nutrient recycling, their interactions with other animals, and their demographic rates. Factors that affect organismal stoichiometry are generally poorly understood, but likely reflect elemental investments in morphological features and life history traits, acting in concert with the environmental availability of elements. We assessed the relative contribution of organismal traits and environmental variability to the stoichiometry of an insectivorous Neotropical stream fish, Rivulus hartii. We characterized the influence of body size, life history phenotype, stage of maturity, and environmental variability on organismal stoichiometry in 6 streams that differ in a broad suite of environmental variables. The elemental composition of R. hartii was variable, and overlapped with the wide range of elemental composition documented across freshwater fish taxa. Average %P composition was ～3.2%(±0.6), average %N～10.7%(±0.9), and average %C～41.7%(±3.1). Streams were the strongest predictor of organismal stoichiometry, and explained up to 18% of the overall variance. This effect appeared to be largely explained by variability in quality of basal resources such as epilithon N:P and benthic organic matter C:N, along with variability in invertebrate standing stocks, an important food source for R. hartii. Organismal traits were weak predictors of organismal stoichiometry in this species, explaining when combined up to 7% of the overall variance in stoichiometry. Body size was significantly and positively correlated with %P, and negatively with N:P, and C:P, and life history phenotype was significantly correlated with %C, %P, C:P and C:N. Our study suggests that spatial variability in elemental availability is more strongly correlated with organismal stoichiometry than organismal traits, and suggests that the stoichiometry of carnivores may not be completely buffered from environmental variability. We discuss the relevance of these findings to ecological stoichiometry theory.     

Species sorting and stoichiometric plasticity control community C:P ratio of first‐order aquatic consumers

Ecological stoichiometry has proven to be invaluable for understanding consumer response to changes in resource quality. Although interactions between trophic levels occur at the community level, most studies focus on single consumer species. In contrast to individual species, communities may deal with trophic mismatch not only through elemental plasticity but also through changes in species composition. Here, we show that a community of first‐order consumers (e.g. zooplankton) is able to adjust its stoichiometry (C:P) in response to experimentally induced changes in resource quality, but only to a limited extent. Furthermore, using the Price equation framework we show the importance of both elemental plasticity and species sorting. These results illustrate the need for a community perspective in ecological stoichiometry, requiring consideration of species‐specific elemental composition, intraspecific elemental plasticity and species turnover.     

Interactions between temperature and nutrients across levels of ecological organization

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits – growth, respiration, body size, and elemental content – that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population‐level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature–nutrient interactions influence processes at the whole‐ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature–nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.     

Deviation from strict homeostasis across multiple trophic levels in an invertebrate consumer assemblage exposed to high chronic phosphorus enrichment in a Neotropical stream

A central tenet of ecological stoichiometry is that consumer elemental composition is relatively independent of food resource nutrient content. Although the P content of some invertebrate consumer taxa can increase as a consequence of P-enriched food resources, little is known about how ecosystem nutrient loading can affect the elemental composition of entire consumer assemblages. Here we examine the potential for P enrichment across invertebrate consumer assemblages in response to chronic high P loading. We measured elemental ratios in invertebrate consumers and basal food resources in a series of streams in lowland Costa Rica that range widely in P levels (2-135 μg l^?1 soluble reactive P). Streams with high P levels receive natural long-term (over millennia) inputs of solute-rich groundwater while low-P streams do not receive these solute-rich groundwater inputs. P content of leaf litter and epilithon increased fourfold across the natural P gradient, exceeding basal resource P content values reported in the literature from other nutrient-rich streams. Invertebrate consumers from the high-P study stream were elevated twofold in P content across multiple taxonomic and functional feeding groups, including predators. Our results strongly support the hypothesis that elevated P content in consumers feeding on P-enriched food resources is a consequence of deviation from strict homeostasis. In contrast to prior studies, we found that between-stream variation in P content of a given taxon greatly exceeded within-stream variation among different taxa, suggesting that environment may be as important as phylogeny in controlling consumer stoichiometry. Relaxing the assumption of strict homeostasis presents challenges and opportunities for advancing our understanding of how nutrient limitation affects consumer growth. Moreover, our findings may provide a window into the future of how chronic anthropogenic nutrient loading can alter stoichiometric relationships in food webs.     

The evolution of ecosystem processes: growth rate and elemental stoichiometry of a key herbivore in temperate and arctic habitats

Understanding the reciprocal interactions between the evolved characteristics of species and the environment in which each species is embedded is a major priority for evolutionary ecology. Here we use the perspective of ecological stoichiometry to test the hypothesis that natural selection on body growth rate affects consumer body stoichiometry. As body elemental composition (nitrogen, phosphorus) of consumers influences nutrient cycling and trophic dynamics in food webs, such differences should also affect biogeochemical processes and trophic dynamics. Consistent with the growth rate hypothesis, body growth rate and phosphorus content of individuals of the Daphnia pulex species complex were lower in Wisconsin compared to Alaska, where the brevity of the growing season places a premium on growth rate. Consistent with stoichiometric theory, we also show that, relative to animals sampled in Wisconsin, animals sampled in Alaska were poor recyclers of P and suffered greater declines in growth when fed low‐quality, P‐deficient food. These results highlight the importance of evolutionary context in establishing the reciprocal relationships between single species and ecosystem processes such as trophic dynamics and consumer‐driven nutrient recycling.     

Thermodynamic constraints on the utility of ecological stoichiometry for explaining global biogeochemical patterns

Carbon and nitrogen cycles are coupled through both stoichiometric requirements for microbial biomass and dissimilatory metabolic processes in which microbes catalyse reduction‐oxidation reactions. Here, we integrate stoichiometric theory and thermodynamic principles to explain the commonly observed trade‐off between high nitrate and high organic carbon concentrations, and the even stronger trade‐off between high nitrate and high ammonium concentrations, across a wide range of aquatic ecosystems. Our results suggest these relationships are the emergent properties of both microbial biomass stoichiometry and the availability of terminal electron acceptors. Because elements with multiple oxidation states (i.e. nitrogen, manganese, iron and sulphur) serve as both nutrients and sources of chemical energy in reduced environments, both assimilative demand and dissimilatory uses determine their concentrations across broad spatial gradients. Conceptual and quantitative models that integrate rather than independently examine thermodynamic, stoichiometric and evolutionary controls on biogeochemical cycling are essential for understanding local to global biogeochemical patterns.     

Linking Microbial and Ecosystem Ecology Using Ecological Stoichiometry: A Synthesis of Conceptual and Empirical Approaches

Currently, one of the biggest challenges in microbial and ecosystem ecology is to develop conceptual models that organize the growing body of information on environmental microbiology into a clear mechanistic framework with a direct link to ecosystem processes. Doing so will enable development of testable hypotheses to better direct future research and increase understanding of key constraints on biogeochemical networks. Although the understanding of phenotypic and genotypic diversity of microorganisms in the environment is rapidly accumulating, how controls on microbial physiology ultimately affect biogeochemical fluxes remains poorly understood. We propose that insight into constraints on biogeochemical cycles can be achieved by a more rigorous evaluation of microbial community biomass composition within the context of ecological stoichiometry. Multiple recent studies have pointed to microbial biomass stoichiometry as an important determinant of when microorganisms retain or recycle mineral nutrients. We identify the relevant cellular components that most likely drive changes in microbial biomass stoichiometry by defining a conceptual model rooted in ecological stoichiometry. More importantly, we show how X-ray microanalysis (XRMA), nanoscale secondary ion mass spectroscopy (NanoSIMS), Raman microspectroscopy, and in situ hybridization techniques (for example, FISH) can be applied in concert to allow for direct empirical evaluation of the proposed conceptual framework. This approach links an important piece of the ecological literature, ecological stoichiometry, with the molecular front of the microbial revolution, in an attempt to provide new insight into how microbial physiology could constrain ecosystem processes.     

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.     

Consumer‐driven nutrient dynamics in freshwater ecosystems: from individuals to ecosystems

The role of animals in modulating nutrient cycling [hereafter, consumer‐driven nutrient dynamics (CND)] has been accepted as an important influence on both community structure and ecosystem function in aquatic systems. Yet there is great variability in the influence of CND across species and ecosystems, and the causes of this variation are not well understood. Here, we review and synthesize the mechanisms behind CND in fresh waters. We reviewed 131 articles on CND published between 1973 and 1 June 2015. The rate of new publications in CND has increased from 1.4 papers per year during 1973–2002 to 7.3 per year during 2003–2015. The majority of investigations are in North America with many concentrating on fish. More recent studies have focused on animal‐mediated nutrient excretion rates relative to nutrient demand and indirect impacts (e.g. decomposition). We identified several mechanisms that influence CND across levels of biological organization. Factors affecting the stoichiometric plasticity of consumers, including body size, feeding history and ontogeny, play an important role in determining the impact of individual consumers on nutrient dynamics and underlie the stoichiometry of CND across time and space. The abiotic characteristics of an ecosystem affect the net impact of consumers on ecosystem processes by influencing consumer metabolic processes (e.g. consumption and excretion/egestion rates), non‐CND supply of nutrients and ecosystem nutrient demand. Furthermore, the transformation and transport of elements by populations and communities of consumers also influences the flow of energy and nutrients across ecosystem boundaries. This review highlights that shifts in community composition or biomass of consumers and eco‐evolutionary underpinnings can have strong effects on the functional role of consumers in ecosystem processes, yet these are relatively unexplored aspects of CND. Future research should evaluate the value of using species traits and abiotic conditions to predict and understand the effects of consumers on ecosystem‐level nutrient dynamics across temporal and spatial scales. Moreover, new work in CND should strive to integrate knowledge from disparate fields of ecology and environmental science, such as physiology and ecosystem ecology, to develop a comprehensive and mechanistic understanding of the functional role of consumers. Comparative and experimental studies that develop testable hypotheses to challenge the current assumptions of CND, including consumer stoichiometric homeostasis, are needed to assess the significance of CND among species and across freshwater ecosystems.     

Elemental stoichiometry of basal resources and benthic macroinvertebrates along a land use gradient in a Great Basin watershed

The effects of land use on the elemental stoichiometry of aquatic organisms have rarely been studied in semi-arid watersheds. In eight semi-arid sub-watersheds differing in land use, we determined which predictor variable(s) best explains the elemental variability in two basal food resources and benthic macroinvertebrates (BMI). The elemental composition of periphyton and seston was best explained by percentage of urban and agricultural areas, forested land and associated differences in SRP, DOC, and stream water N:P ratios. In contrast, consumer elemental stoichiometry was related to taxonomic identity and feeding mode. Elemental imbalances were higher for collector-gatherer than for scraper and collector-filterer. However, high spatial and temporal variability in the elemental composition of basal food resources obscured clear spatial patterns of imbalances between nutrient-poor upstream and nutrient-rich downstream sites. Results from this study suggest that land use can affect BMI due to alteration in stoichiometry of their food resources. However, taxonomy and allometry must be taken into account to better understand spatial and temporal changes in the elemental composition of BMI. Our results indicate the importance of considering multiple effects to accurately assess land use effects on producer and consumer stoichiometry, particularly the in highly variable Great Basin watersheds.     

Using ecological stoichiometry to understand and predict infectious diseases

A key characteristic of host–parasite interactions is the theft of host nutrients by the parasite, yet we lack a general framework for understanding and predicting the interplay of host and parasite nutrition that applies across biological levels of organization. The elemental nutrients (C, N, P, Fe, etc.), and ecological stoichiometry provide a framework for understanding host–parasite interactions and their relation to ecosystem functioning. Here we use the ecological stoichiometry framework to develop hypotheses and predictions regarding the relationship between elemental nutrients and host–parasite interactions. We predict that a suite of host and parasite traits, stoichiometric homeostasis, host diet stoichiometry, and biogeochemical cycling are related to disease dynamics, host immunity and resistance, and bacterial growth form determination. We show that ecological stoichiometry is capable of expanding our understanding of host–parasite interactions, and complementing other approaches such as population and community ecology, and molecular biology, for studying infectious diseases.     

Temporal changes in soil C‐N‐P stoichiometry over the past 60 years across subtropical China

Controlled experiments have shown that global changes decouple the biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P), resulting in shifting stoichiometry that lies at the core of ecosystem functioning. However, the response of soil stoichiometry to global changes in natural ecosystems with different soil depths, vegetation types, and climate gradients remains poorly understood. Based on 2,736 observations along soil profiles of 0–150 cm depth from 1955 to 2016, we evaluated the temporal changes in soil C‐N‐P stoichiometry across subtropical China, where soils are P‐impoverished, with diverse vegetation, soil, and parent material types and a wide range of climate gradients. We found a significant overall increase in soil total C concentration and a decrease in soil total P concentration, resulting in increasing soil C:P and N:P ratios during the past 60 years across all soil depths. Although average soil N concentration did not change, soil C:N increased in topsoil while decreasing in deeper soil. The temporal trends in soil C‐N‐P stoichiometry differed among vegetation, soil, parent material types, and spatial climate variations, with significantly increased C:P and N:P ratios for evergreen broadleaf forest and highly weathered Ultisols, and more pronounced temporal changes in soil C:N, N:P, and C:P ratios at low elevations. Our sensitivity analysis suggests that the temporal changes in soil stoichiometry resulted from elevated N deposition, rising atmospheric CO₂ concentration and regional warming. Our findings revealed that the responses of soil C‐N‐P and stoichiometry to long‐term global changes have occurred across the whole soil depth in subtropical China and the magnitudes of the changes in soil stoichiometry are dependent on vegetation types, soil types, and spatial climate variations.     

生态系统模拟模型的研究进展

从四个方面概述了生态系统模拟模型的发展现状：1)个体及种群,种群动态模型主要模拟在一个生境中单个种的动、植物个体出生或发芽、成长及其死亡过程,还有种内竞争和种间相互作用,主要分析生境中生物之间的相互作用。主要概述了林窗模型和土壤一植物一大气系统模型。2)群落与生态系统,概述了生态系统生产力模型、生物地球化学循环模型及演替模型。主要模拟植物种类在整个生态系统发展过程中的变化,以及植被类型的转变和相关的生物地球化学循环过程的改变,从而反映生物群落对气候变化的响应。3)景观生态系统,景观动态研究包含了时空两个方面的动态变化,一般可分为随机景观模型和基于过程的景观模型。随机模型用于模拟群落格局在演替过程中的动态变化等,基于过程的景观模型深入研究组成景观的各生态系统的空间结构。4)生物圈与地球生态系统,基于过程的陆地生物地球化学模式被用来研究自然生态系统中碳和其它矿物营养物质的潜在通量和蓄积量,较为流行的模式有陆地生态系统模式TEM、CENTURY、法兰克福生物圈模式FBM、Biome-BGC、卡内基-埃姆斯-斯坦福方法CASA等。这些模式己被用于估算自然生态系统对大气CO2加倍及相关气候变化在区域和全球尺度的平衡响应。最后,结合实际工作展望了生态系统模拟模型在各方面的发展方向。     

Meta-ecosystems: a theoretical framework for a spatial ecosystem ecology

This contribution proposes the meta‐ecosystem concept as a natural extension of the metapopulation and metacommunity concepts. A meta‐ecosystem is defined as a set of ecosystems connected by spatial flows of energy, materials and organisms across ecosystem boundaries. This concept provides a powerful theoretical tool to understand the emergent properties that arise from spatial coupling of local ecosystems, such as global source–sink constraints, diversity–productivity patterns, stabilization of ecosystem processes and indirect interactions at landscape or regional scales. The meta‐ecosystem perspective thereby has the potential to integrate the perspectives of community and landscape ecology, to provide novel fundamental insights into the dynamics and functioning of ecosystems from local to global scales, and to increase our ability to predict the consequences of land‐use changes on biodiversity and the provision of ecosystem services to human societies.