食物链长度理论研究进展

食物链长度（Food chain length,FCL）是生态系统中最重要的特点之一,它通过改变生物间的营养关系,影响着生物多样性,群落的结构以及稳定性;它是反映食物网物质转换与能量传递的综合指数,食物链及其动态特征是生态学许多重要理论的基础,食物链长度理论的研究进展,推动了人们对水域生态系统中生物和非生物相互作用的理解。回顾了食物链长度的3种度量方法及其详细的计算方法,在此基础上简述了各方法的特点。综述了食物链长度的决定因素的4种假说（资源可利用性假说、生产力空间假说、生态系统大小假说、动态稳定性假说）及其交互作用,重点总结了湖泊食物链长度的空间格局与决定因素的研究进展。最后,食物链长度研究展望,包括食物链长度决定因子研究存在的问题及发展方向的总结,以及在在水域生态学中的应用的研究进展,例如食物链长度在指示污染物的生物富集中的研究进展、食物链食物链长度在指导生物操作、以及在食物链长度在对气候变化响应方面的研究进展等等。     

Refuge increases food chain length: modeled impacts of littoral structure in lake food webs

Food chain length (FCL) represents a fundamental metric within ecology because it has implications for ecosystem function and responses to environmental change. Omnivory between linked food chains situated within large ecosystems can increase FCL, whereas overlap of food chains within small or spatially compressed ecosystems is generally thought to decrease FCL. Yet FCL varies widely in small ecosystems and the mechanisms underlying determinants of FCL in these systems is unclear. In small shallow lakes, littoral structure is a predictor of FCL but it is unclear whether this is due to productivity or refuge mechanisms. Here we provide evidence, using consumer resource food web modules parameterized with empirical data, that refuge in spatially compressed ecosystems has the ability on its own to increase the trophic position of top predators by increasing the biomass of top and intermediate predators across a range of common food web module structures. Our results suggest that refuge is an important driver of FCL in small ecosystems, which has implications for determining responses of these systems to environmental change.     

Resource availability and ecosystem size predict food-chain length in pond ecosystems

Food-chain length (FCL) in ecosystems has been studied extensively, and numerous hypotheses to predict FCL, productivity, ecosystem size, and productive space have been proposed. For example, the productivity hypothesis suggests that resource availability limits FCL, whereas the productive-space hypothesis predicts that per-unit-size resource availability and ecosystem size equally limit FCL. However, previous studies have only measured total productivity to test FCL and have never tested the impact of resource availability within a system on FCL. Therefore, we estimated FCL in 15 ponds using stable isotope techniques to test the most common hypotheses for predicting FCL. We also measured total productivity, pond volume (ecosystem size), and edible microalgal carbon (resource availability) in each pond. We found that productive-space (edible carbon+pond volume) was the best model to predict FCL, and FCL was significantly correlated with edible carbon and pond volume. However, factors such as total productivity did not directly predict FCL of the ponds. Therefore, our results suggest that both resource availability and ecosystem size predict FCL in pond ecosystems and play significant roles in maintaining longer FCLs. Furthermore, the productive-space hypothesis appears to be particularly important for determining the FCL of ponds.     

Resource productivity and availability impacts for food-chain length

Ecologists have focused on food-chain length (FCL) for the past eight decades as an index of food-web structure. Here, I review the hypotheses determining FCL with a focus on resource productivity and availability effects on FCL. First, I introduce the mainstream hypotheses to explain FCL variations: productivity, ecosystem size, and disturbance. For the existing productivity and productive space hypotheses, I stress the importance of using resource availability to estimate the productivity effect on FCL. Using a FCL dataset from 15 ponds, I tested the resource stoichiometry effect on FCL for ponds with between carbon:nitrogen ratio of primary producers and FCL. Moreover, I provide a perspective for studying resource availability and stoichiometry effects on FCL and of the alternative hypotheses of productive-space and ecosystem size. Finally, I suggest the future directions of the FCL study: a resource subsidy and climate change effects on FCL and food-web structure.     

Predicting Plant Diversity Response to Disturbance: Applicability of the Intermediate Disturbance Hypothesis and Mass Ratio Hypothesis

Predicting the relationships between disturbance, biodiversity and productivity of ecosystems continue to preoccupy ecologists and resource managers. Two hypotheses underpin many of the discussions. The Intermediate Disturbance Hypothesis (IDH), which proposes that biodiversity peaks at intermediate levels of disturbance, is often extended to predict that productivity follows the same response pattern. The Mass Ratio Hypothesis (MRH) proposes that the biological traits of the dominant species are the critical drivers of ecosystem function (e.g., productivity) and that these species increase in biomass rapidly after disturbance then stabilize. As a consequence, species diversity first peaks then declines after disturbance as a few species dominate the site. Both provide a conceptual link among disturbance, species diversity and productivity (an index of ecosystem function). We assessed the current state of empirical support for these two hypotheses with a literature survey and determined if their conformance is related to ecosystem type or site productivity. Conformance of IDH reported in past reviews (considering all ecosystems) ranged from 16 to 21%. This contrasts with our finding that in terrestrial ecosystems conformance to IDH was 46% (22 of 48 studies), 17% studies reported non-compliance, and 23% reported inconclusive results. Most studies explained their results with respect to IDH or MRH. Only two studies were specifically designed to test the validity of IDH or MRH. We conclude that (i) the IDH is mostly applicable to predict species diversity response to disturbance in upland sites of medium to high productivity and the MRH is applicable to organic sites of low productivity; (ii) there is a critical need for more studies specifically designed to test these hypotheses in natural ecosystems using common protocols; and (iii) enhanced understanding of these models will add value in refining management policies and in the selection of meaningful diversity indicators of sustainability.     

Temperature‐size responses alter food chain persistence across environmental gradients

Body‐size reduction is a ubiquitous response to global warming alongside changes in species phenology and distributions. However, ecological consequences of temperature‐size (TS) responses for community persistence under environmental change remain largely unexplored. Here, we investigated the interactive effects of warming, enrichment, community size structure and TS responses on a three‐species food chain using a temperature‐dependent model with empirical parameterisation. We found that TS responses often increase community persistence, mainly by modifying consumer‐resource size ratios and thereby altering interaction strengths and energetic efficiencies. However, the sign and magnitude of these effects vary with warming and enrichment levels, TS responses of constituent species, and community size structure. We predict that the consequences of TS responses are stronger in aquatic than in terrestrial ecosystems, especially when species show different TS responses. We conclude that considering the links between phenotypic plasticity, environmental drivers and species interactions is crucial to better predict global change impacts on ecosystem diversity and stability.     

Environmental determinants of food-chain length: a meta-analysis

Food-chain length is an important character of ecological communities that affects many of their functional aspects. Recently, an increasing number of studies have tested the effects of productivity, disturbance, or ecosystem size on food-chain length in a variety of natural systems. Here we conduct a formal meta-analysis to summarize findings from these empirical studies. We found significant positive mean effects of productivity and ecosystem size but no significant mean effect of disturbance on food-chain length. The strength of mean effect sizes was not significantly different between productivity and ecosystem size. These results lend general support to previous theories predicting the effect of productivity and ecosystem size, but fail to support the prediction that disturbance shortens food chains. In addition, our meta-analysis found that the effect sizes of primary studies were significantly heterogeneous for ecosystem size and disturbance, but not for productivity. This pattern might reflect that ecosystem size and disturbance can affect food-chain length through multiple different mechanisms, while productivity influences food-chain length in a simple manner through energy limitation.     

Will a large complex system be productive?

While the relationship between food web complexity and stability has been well documented, how complexity affects productivity remains elusive. In this study, we combine food web theory and a data set of 149 aquatic food webs to investigate the effect of complexity (i.e. species richness, connectance, and average interaction strength) on ecosystem productivity. We find that more complex ecosystems tend to be more productive, although different facets of complexity have contrasting effects. A higher species richness and/or average interaction strength increases productivity, whereas a higher connectance often decreases it. These patterns hold not only between realized complexity and productivity, but also characterize responses of productivity to simulated declines of complexity. Our model also predicts a negative association between productivity and stability along gradients of complexity. Empirical analyses support our predictions on positive complexity-productivity relationships and negative productivity-stability relationships. Our study provides a step forward towards reconciling ecosystem complexity, productivity and stability.     

Global patterns of aquatic food chain length

Food chain length is a fundamental ecosystem property, and plays a central role in determining ecosystem functioning. Recent advances in the field of stable isotope ecology allow the estimation of food chain length (FCL) from stable nitrogen isotope (δ¹⁵N) data. We conducted a global literature synthesis and estimated FCL for 219 lake, stream, and marine ecosystems. Streams had shorter food chains (∼3.5 trophic levels) than marine and lake ecosystems (∼4.0 trophic levels). In marine systems, inclusion of marine mammals increased FCL by 2/3 of a trophic level. For each ecosystem type, estimates of FCL were normally distributed and spanned two full trophic levels. Comparison with published connectance food webs revealed similar mean FCL values, though stable isotope-derived FCL estimates were less variable. At the global scale, FCL showed weak or no relationships with ecosystem size, mean annual air temperature, or latitude. Our study highlights the utility of stable isotopes for quantifying among-system food web variability, and the application of this approach for assessing global-scale patterns of food chain length.     

食物链长度远因与近因研究进展综述

食物链长度是生态系统的基本属性,其变化决定着群落结构和生态系统功能。稳定同位素分析技术的进步推进了生态系统中食物链长度决定因子相关研究的开展。尽管近期的研究证明了食物链长度与资源可利用性、生态系统大小、干扰等远因之间的关系,但是对于食物网内部结构变化这一近因对食物链长度的影响作用关注较少。综述了边界明确和开放类型淡水生态系统中食物链长度的相关研究进展;探讨了远因和近因机制在决定食物链长度中的作用;给出了判断不同层次和尺度上决定食物链长度机制的概念框架;为今后更好的开展不同生态系统间食物链长度的比较研究提出了建议。     

An assessment of natural and human disturbance effects on Mexican ecosystems: current trends and research gaps

Mexico harbors more than 10% of the planet’s endemic species. However, the integrity and biodiversity of many ecosystems is experiencing rapid transformation under the influence of a wide array of human and natural disturbances. In order to disentangle the effects of human and natural disturbance regimes at different spatial and temporal scales, we selected six terrestrial (temperate montane forests, montane cloud forests, tropical rain forests, tropical semi-deciduous forests, tropical dry forests, and deserts) and four aquatic (coral reefs, mangrove forests, kelp forests and saline lakes) ecosystems. We used semi-quantitative statistical methods to assess (1) the most important agents of disturbance affecting the ecosystems, (2) the vulnerability of each ecosystem to anthropogenic and natural disturbance, and (3) the differences in ecosystem disturbance regimes and their resilience. Our analysis indicates a significant variation in ecological responses, recovery capacity, and resilience among ecosystems. The constant and widespread presence of human impacts on both terrestrial and aquatic ecosystems is reflected either in reduced area coverage for most systems, or reduced productivity and biodiversity, particularly in the case of fragile ecosystems (e.g., rain forests, coral reefs). In all cases, the interaction between historical human impacts and episodic high intensity natural disturbance (e.g., hurricanes, fires) has triggered a reduction in species diversity and induced significant changes in habitat distribution or species dominance. The lack of monitoring programs assessing before/after effects of major disturbances in Mexico is one of the major limitations to quantifying the commonalities and differences of disturbance effects on ecosystem properties.     

Species richness facilitates ecosystem resilience in aquatic food webs

1. Many studies indicate that biodiversity in ecosystems affects stability, either by promoting temporal stability of ecosystem attributes or by enhancing ecosystem resistance and resilience to perturbation. The effects on temporal stability are reasonably well understood and documented but effects on resistance and resilience are not. 2. Here, we report results from an aquatic mesocosm experiment in which we manipulated the species richness and composition of aquatic food webs (macrophytes, macro‐herbivores and invertebrate predators), imposed a pulse disturbance (acidification), and monitored the resistance (initial response) and resilience (recovery) of ecosystem productivity and respiration. 3. We found that species‐rich macroinvertebrate communities had higher resilience of whole‐ecosystem respiration, but were not more resistant to perturbations. We also found that resilience and resistance were unaffected by species composition, despite the strong role composition is known to play in determining mean levels of function in these communities. 4. Biodiversity’s effects on resilience were probably mediated through complex pathways affecting phytoplankton and microbial communities (e.g. via changes in nutrient regeneration, grazing or compositional changes) rather than through simpler effects (e.g. insurance effects, enhanced facilitation) although these simpler mechanisms probably played minor roles in enhancing respiration resilience. 5. Current mechanisms for understanding biodiversity’s effects on ecosystem stability have been developed primarily in the context of single‐trophic level communities. These mechanisms may be overly simplistic for understanding the consequences of species richness on ecosystem stability in complex, multi‐trophic food webs where additional factors such as indirect effects and highly variable life‐history traits of species may also be important.     

Effects of plant diversity,community composition and environmental parameters on productivity in montane European grasslands

In the past years, a number of studies have used experimental plant communities to test if biodiversity influences ecosystem functioning such as productivity. It has been argued, however, that the results achieved in experimental studies may have little predictive value for species loss in natural ecosystems. Studies in natural ecosystems have been equivocal, mainly because in natural ecosystems differences in diversity are often confounded with differences in land use history or abiotic parameters. In this study, we investigated the effect of plant diversity on ecosystem functioning in semi-natural grasslands. In an area of 10×20 km, we selected 78 sites and tested the effects of various measures of diversity and plant community composition on productivity. We separated the effects of plant diversity on ecosystem functioning from potentially confounding effects of community composition, management or environmental parameters, using multivariate statistical analyses. In the investigated grasslands, simple measures of biodiversity were insignificant predictors of productivity. However, plant community composition explained productivity very well (R²=0.31) and was a better predictor than environmental variables (soil and site characteristics) or management regime. Thus, complex measures such as community composition and structure are important drivers for ecosystem functions in semi-natural grasslands. Furthermore, our data show that it is difficult to extrapolate results from experimental studies to semi-natural ecosystems, although there is a need to investigate natural ecosystems to fully understand the relationship of biodiversity and ecosystem functioning.     

Robustness of size–structure across ecological networks in pelagic systems

The study of biomass size distributions has become an important tool for addressing aquatic ecosystem complexity and the consequences of anthropogenic disturbances. However, it remains unclear how changes in pelagic food web topology affect the biomass size–structure. Employing a dynamic multispecies bioenergetic consumer-resource model, we simulated biomass trajectories over time in 10,000 virtual networks of varying topology to address which food web properties are important in determining size–structure in pelagic systems. The slopes of the normalized biomass size spectra (NBSS) and Pareto’s shape parameter (γ) of our modeled communities are consistent with theoretically expected values for steady-state systems and empirical values reported for several aquatic ecosystems. We found that the main drivers of the NBSS slope and Pareto’s γ were the slope of the relationship between body mass and trophic level, the maximum trophic level of the food web, and the stability of total community biomass. Our analyses showed a clear conservative trend in pelagic community size–structure as demonstrated by the robustness of the NBSS slope and Pareto’s γ against most of the topological changes in virtual networks. Nevertheless, these analyses also caution that major disturbances in large-bodied or top-trophic level individuals may disrupt this stable pattern.     

Effects of productivity, disturbance, and ecosystem size on food-chain length: insights from a metacommunity model of intraguild predation

Traditionally, productivity and disturbance have been hypothesized as important determinants of food-chain length. More recently, growing empirical evidence suggests a strong role of ecosystem size. To theoretically explore the effects of basal productivity, disturbance, and ecosystem size on food-chain length, we develop and analyze a metacommunity model of intraguild predation (IGP). The model finds that, when local IGP is weak, increasing basal productivity, weakening disturbance, and increasing ecosystem size will generally increase food-chain length. When local IGP is strong, by contrast, increasing basal productivity or weakening disturbance favors intraguild predators and hinders the coexistence of intraguild predators and intraguild prey, limiting food-chain length. In contrast, increasing ecosystem size can promote coexistence even when local IGP is strong, increasing food-chain length through inserting intraguild prey and changing the degree of omnivory by intraguild predators. Intraguild prey needs to be the superior colonizer to intraguild predators for this to occur. We discuss that these theoretical predictions appear consistent with empirical patterns.     

Crossing habitat boundaries: coupling dynamics of ecosystems through complex life cycles

Ecosystems are often indirectly connected through consumers with complex life cycles (CLC), in which different life stages inhabit different ecosystems. Using a structured consumer resource model that accounts for the independent effects of two resources on consumer growth and reproductive rates, we show that such indirect connections between ecosystems can result in alternative stable states characterized by adult-dominated and juvenile-dominated consumer populations. As a consequence, gradual changes in ecosystem productivity or mortality rates of the consumer can lead to dramatic and abrupt regime shifts across different ecosystems, hysteresis and counterintuitive changes in the consumer abundances. Whether these counter intuitive or abrupt responses occur depend on the relative productivity of both habitats and which consumer life-stage inhabits the manipulated ecosystem. These results demonstrate the strong yet complex interactions between ecosystems coupled through consumers with CLC and the need to think across ecosystems to reliably predict the consequences of natural or anthropogenic changes.     

气候变化、火干扰与生态系统生产力

 综述了气候变化、火干扰与生态系统生产力之间的相互作用关系以及目前相关的研究进展。侧重介绍了气候变化与火干扰之间的相互作用关系以及火干扰对生态系统生产力的影响。气候变化通过作用于可燃物质数量、湿度和火灾天气来影响火干扰的发生频率和强度,而火干扰过程释放大量温室气体和烟尘物质反过来也会对气候变化产生影响。另外,火干扰过程改变了火烧迹地的土壤生物地球化学性质、养分循环和分配以及大气组成,进而对生态系统对CO2的吸收能力产生影响。正确理解三者之间的逻辑关系,对于我们有效地利用火管理提高区域生态系统碳吸收,减少碳排放,减缓全球变化速率,都具有重要的指导意义 。     

All wet or dried up? Real differences between aquatic and terrestrial food webs

Ecologists have greatly advanced our understanding of the processes that regulate trophic structure and dynamics in ecosystems. However, the causes of systematic variation among ecosystems remain controversial and poorly elucidated. Contrasts between aquatic and terrestrial ecosystems in particular have inspired much speculation, but only recent empirical quantification. Here, we review evidence for systematic differences in energy flow and biomass partitioning between producers and herbivores, detritus and decomposers, and higher trophic levels. The magnitudes of different trophic pathways vary considerably, with less herbivory, more decomposers and more detrital accumulation on land. Aquatic-terrestrial differences are consistent across the global range of primary productivity, indicating that structural contrasts between the two systems are preserved despite large variation in energy input. We argue that variable selective forces drive differences in plant allocation patterns in aquatic and terrestrial environments that propagate upward to shape food webs. The small size and lack of structural tissues in phytoplankton mean that aquatic primary producers achieve faster growth rates and are more nutritious to heterotrophs than their terrestrial counterparts. Plankton food webs are also strongly size-structured, while size and trophic position are less strongly correlated in most terrestrial (and many benthic) habitats. The available data indicate that contrasts between aquatic and terrestrial food webs are driven primarily by the growth rate, size and nutritional quality of autotrophs. Differences in food-web architecture (food chain length, the prevalence of omnivory, specialization or anti-predator defences) may arise as a consequence of systematic variation in the character of the producer community.     

Flow regulation increases food‐chain length through omnivory mechanisms in a Mediterranean river network

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