Chemical information transfer in freshwater plankton

The structure of aquatic ecosystems is determined by complex interactions among individual organisms at different trophic levels. Although our basic understanding of how top-down and bottom-up processes interact to determine food-web dynamics has advanced, we still lack insights into how complex interactions and feedbacks affect the dynamics and structure of food webs. It is now becoming increasingly clear that, in addition to energy transfer from one trophic level to the other, there is exchange of information between these levels facilitated by the release of infochemicals by the organisms. There is evidence from recent studies that the exchange of chemical information in freshwater ecosystems is likely to play a decisive role in shaping structure and functioning of these systems. Chemical communication among freshwater organisms mediates many aspects of both predation and interspecific competition, which play key roles in determining community structure and ecosystem functioning. For example, consumer-induced defences in phytoplankton and zooplankton include modifications in the characteristics relating to life history, behaviour, morphology and biochemistry. These inducible defences affect trophic interactions by altering predator feeding rates through changes in attack rate or handling time, or both. Also host-specific fungal parasitism in phytoplankton is probably controlled by infochemicals. The motile fungi recognise their host by host-secreted compounds. Until now models describing the functioning of ecosystems mainly considered flows of biomass and energy. Integration of new knowledge about the role of chemical communication in these models may be one of the aims of ecological informatics. In this chapter I discuss how infochemicals may affect the dynamics and structure of planktonic food webs.     

Effects of trophic skewing of species richness on ecosystem functioning in a diverse marine community

Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.     

Species loss and the structure and functioning of multitrophic aquatic systems

Experiments and theory in single trophic level systems dominate biodiversity and ecosystem functioning research and recent debates. All natural ecosystems contain communities with multiple trophic levels, however, and this can have important effects on ecosystem structure and functioning. Furthermore, many experiments compare assembled communities, rather than examining loss of species directly. We identify three questions around which to organise an investigation of how species loss affects the structure and functioning of multitrophic systems. 1) What is the distribution of species richness among trophic levels; 2) from which trophic levels are species most often lost; and 3) does loss of species from different trophic levels influence ecosystem functioning differently? Our analyses show that: 1) Relatively few high‐quality data are available concerning the distribution of species richness among trophic levels. A new data‐set provides evidence of a decrease in species richness as trophic height increases. 2) Multiple lines of evidence indicate that species are lost from higher trophic levels more frequently than lower trophic levels. 3) A theoretical model suggests that both the structure of food webs (occurrence of omnivory and the distribution of species richness among trophic levels) and the trophic level from which species are lost determines the impact of species loss on ecosystem functioning, which can even vary in the sign of the effect. These results indicate that, at least for aquatic systems, models of single trophic level ecosystems are insufficient for understanding the functional consequences of extinctions. Knowledge is required of food web structure, which species are likely to be lost, and also whether cascading extinctions will occur.     

水生生态系统食物网复杂性与多样性的关系

探索食物网的复杂结构是生态学的中心问题之一。基于构建的黄河口海草床食物网并耦合实际食物网的数据集,整理了包含河口、湖泊、海洋和河流四种水生生态系统类型的48个实际食物网案例。以食物网的节点数反映食物网多样性,物种之间的营养链接数、链接密度和连通度来表示食物网的复杂性,采用营养缩尺模型描述水生生态系统食物网的复杂性特征与节点数的普适性规律。结果表明：所涉及的48个水生生态系统食物网的多样性和复杂性跨度较大,其中,节点数的分布范围为4-124,链接数为3-1830,链接密度为0.75-15.71,连通度为0.06-0.25。不同类型水生生态系统间的连通度存在显著性差异（P=0.01）,节点数、链接数、链接密度不存在显著性差异。各类型生态系统的食物网链接数、链接密度均随节点数的增加而增加（R²=0.92,P<0.001和R²=0.82,P<0.001）。湖泊生态系统的连通度随节点数的变化不明显,围绕在0.20附近;而其他3种类型生态系统的食物网连通度随节点数的增加而降低（R²=0.06-0.41,P<0.001）。对全球尺度的水生食物网多样性和复杂性的定量化研究对于提升对食物网的复杂结构的科学认识,从系统尺度探究多样性和复杂性的关系提供数据支撑。     

Trophic roles of tadpoles in tropical Australian streams

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Trophic networks: How do theories link ecosystem structure and functioning to stability properties? A review

In the context of present global changes, interest in understanding how systems respond to anthropogenic environmental pressures and stress has increased. Indices that characterize ecosystem state are helpful tools for the interpretation of ecosystem responses. The central question is how to link these responses to ecosystem structure and functioning and to quantify ecosystem persistence, resistance or resilience. Quantification and characterization of trophic networks by ecological network analysis (ENA) indices is proceeding rapidly, especially in the field of coastal ecology. In this contribution, we review several theories that relate ecosystem structure and function to stability. The structure and functioning of ecosystems change during the maturation of ecosystems. In the first section, the maturation of ecosystems is described using thermodynamics. In the second and third parts of this paper, we define some concepts for analysing structure and functioning of food webs and discuss their relation to stability. In the last section, we describe three ENA indices and their link to stability. We demonstrate that ENA provides powerful tools for describing local stability, combining quantitative and qualitative concepts. However, it remains incomplete for describing real conservation cases that combine local and global stability.     

Pyramids of species richness: the determinants and distribution of species diversity across trophic levels

How species richness is distributed across trophic levels determines several dimensions of ecosystem functioning, including herbivory, predation, and decomposition rates. We perform a meta‐analysis of 72 large published food webs to investigate their trophic diversity structure and possible endogenous, exogenous, and methodological causal variables. Consistent with classic theory, we found that published food webs can generally be described as ‘pyramids of species richness’. The food webs were more predator‐poor, prey‐rich and hierarchical than is expected by chance or by the niche or cascade models. The trophic species richness distribution also depended on centrality, latitude, ecosystem‐type and methodological bias. Although trophic diversity structure is generally pyramidal, under many conditions the structure is consistently uniform or inverse‐pyramidal. Our meta‐analysis adds nuance to classic assumptions about food web structure: diversity decreases with trophic level, but not under all conditions, and the decrease may be scale‐dependent. Synthesis The distribution of species richness across trophic levels has not been evaluated in recent decades, despite improvement in food web resolution and the relevance of biodiversity distribution to ecosystem function. Our meta‐analysis of 72 large, recent food webs, illustrates that published food webs can generally be described as basal‐rich, top‐poor ‘pyramids of species richness’, consistent with classic theory. Although trophic diversity structure is generally pyramidal, under some environmental and ecological conditions the structure is uniform or inverse‐pyramidal. Our meta‐analysis confirms classic theory about food web structure, while adding nuance by describing conditions under which classic pyramid structure is not observed.     

Food-web structure of seagrass communities across different spatial scales and human impacts

Seagrass beds provide important habitat for a wide range of marine species but are threatened by multiple human impacts in coastal waters. Although seagrass communities have been well-studied in the field, a quantification of their food-web structure and functioning, and how these change across space and human impacts has been lacking. Motivated by extensive field surveys and literature information, we analyzed the structural features of food webs associated with Zostera marina across 16 study sites in 3 provinces in Atlantic Canada. Our goals were to (i) quantify differences in food-web structure across local and regional scales and human impacts, (ii) assess the robustness of seagrass webs to simulated species loss, and (iii) compare food-web structure in temperate Atlantic seagrass beds with those of other aquatic ecosystems. We constructed individual food webs for each study site and cumulative webs for each province and the entire region based on presence/absence of species, and calculated 16 structural properties for each web. Our results indicate that food-web structure was similar among low impact sites across regions. With increasing human impacts associated with eutrophication, however, food-web structure show evidence of degradation as indicated by fewer trophic groups, lower maximum trophic level of the highest top predator, fewer trophic links connecting top to basal species, higher fractions of herbivores and intermediate consumers, and higher number of prey per species. These structural changes translate into functional changes with impacted sites being less robust to simulated species loss. Temperate Atlantic seagrass webs are similar to a tropical seagrass web, yet differed from other aquatic webs, suggesting consistent food-web characteristics across seagrass ecosystems in different regions. Our study illustrates that food-web structure and functioning of seagrass habitats change with human impacts and that the spatial scale of food-web analysis is critical for determining results.     

Food web structure in Mediterranean streams: exploring stabilizing forces in these ecosystems

We constructed the food webs of six Mediterranean streams in order to determine ecological generalities derived from analysis of their structure and to explore stabilizing forces within these ecosystems. Fish, macroinvertebrates, primary producers and detritus are the components of the studied food webs. Analysis focused on a suite of food web properties that describe species’ trophic habits, linkage complexity and food chains. A great structural similarity was found in analyzed food webs; we therefore suggest average values for the structural properties of Mediterranean stream food webs. Percentage of omnivorous species was positively correlated with connectance, and there was a predominance of intermediate trophic level species that had established simple links with detritus. In short, our results suggest that omnivory and the weak interactions of detritivores have a stabilizing role in these food webs.     

食物网结构与功能: 理论进展与展望

食物网刻画了物种间通过捕食而形成的复杂网络关系。阐明食物网结构与功能之间的关系, 既是生态学的基本理论问题, 也是预测全球变化背景下生态系统响应的重要依据。早期关于食物网结构与功能的研究往往是分离的, 或是基于食物链等的简单网络模型, 而近期研究基于复杂食物网模型取得了重要理论进展。本文综述了食物网研究的理论方法和近期进展, 特别介绍了复杂食物网中的结构、多样性和功能的度量指标、结构-多样性-功能之间的关系以及全球变化对食物网结构与功能的影响。本文最后对未来的一些研究方向进行了展望, 包括与功能性状和化学计量学的整合、食物网与其他网络类型的整合以及拓展食物网研究的空间和时间尺度。     

Biodiversity and ecosystem functioning in food webs: the vertical diversity hypothesis

One challenge in merging community and ecosystem ecology is to integrate the complexity of natural multitrophic communities into concepts of ecosystem functioning. Here, we combine food‐web and allometry theories to demonstrate that primary production, as measured by the total nutrient uptake of the multitrophic community, is determined by vertical diversity (i.e. food web's maximum trophic level) and structure (i.e. distributions of species and their abundances and metabolic rates across trophic levels). In natural ecosystems, the community size distribution determines all these vertical patterns and thus the total nutrient uptake. Our model suggests a vertical diversity hypothesis (VDH) for ecosystem functioning in complex food webs. It predicts that, under a given nutrient supply, the total nutrient uptake increases exponentially with the maximum trophic level in the food web and it increases with its maximum body size according to a power law. The VDH highlights the effect of top–down regulation on plant nutrient uptake, which complements traditional paradigms that emphasised the bottom–up effect of nutrient supply on vertical diversity. We conclude that the VDH contributes to a synthetic framework for understanding the relationship between vertical diversity and ecosystem functioning in food webs and predicting the impacts of global changes on multitrophic ecosystems.     

Biodiversity, ecosystem functioning and food webs in fresh waters: assembling the jigsaw puzzle

1. Dramatic advances have been made recently in the study of biodiversity–ecosystem functioning (B-EF) relations and food web ecology. These fields are now starting to converge, and this fusion has the potential to improve our understanding of how environmental stressors modulate ecosystem processes and the supply of 'goods and services'.
2. Food web structure and dynamics can exert particularly strong influences on B-EF relations in fresh waters, as consumer–resource interactions (e.g. trophic cascades) are often more important than horizontal interactions within trophic levels. For instance, many freshwater food webs are size structured, with large organisms tending to occupy the higher trophic levels and often exerting powerful effects on ecosystem processes. However, because they are also vulnerable to perturbations, non-random losses of these large taxa can alter both food web structure and ecosystem functioning profoundly.
3. Recently, the focus of food web research has shifted away from exploring patterns, towards developing an understanding of processes (e.g. quantifying fluxes of individuals, biomass, energy, nutrients) and how the two interact. Many of the best-characterized food webs are from fresh waters, and these ecosystems are now being used to address some of the shortcomings of earlier B-EF studies. I have identified several key gaps in our current knowledge and highlighted potentially fruitful avenues of future B-EF and food web research.
4. A major challenge for this newly emerging research is to place it within a unified theoretical framework. The application of metabolic theory and ecological stoichiometry may help to achieve this goal by considering biological systems within the constraints imposed upon them by physical and chemical laws.     

Compensation masks trophic cascades in complex food webs

Ecological networks, or food webs, describe the feeding relationships between interacting species within an ecosystem. Understanding how the complexity of these networks influences their response to changing top-down control is a central challenge in ecology. Here, we provide a model-based investigation of trophic cascades — an oft-studied ecological phenomenon that occurs when changes in the biomass of top predators indirectly effect changes in the biomass of primary producers — in complex food webs that are representative of the structure of real ecosystems. Our results reveal that strong cascades occur primarily in low richness and weakly connected food webs, a result in agreement with some prior predictions. The primary mechanism underlying weak or absent cascades was a strong compensatory response; in most webs, predators induced large population level cascades that were masked by changes in the opposite direction by other species in the same trophic guild. Thus, the search for a general theory of trophic cascades in food webs should focus on uncovering features of real ecosystems that promote biomass compensation within functional guilds or trophic levels.     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

Loss of Rare Fish Species from Tropical Floodplain Food Webs Affects Community Structure and Ecosystem Multifunctionality in a Mesocosm Experiment

Experiments with realistic scenarios of species loss from multitrophic ecosystems may improve insight into how biodiversity affects ecosystem functioning. Using 1000 L mesocoms, we examined effects of nonrandom species loss on community structure and ecosystem functioning of experimental food webs based on multitrophic tropical floodplain lagoon ecosystems. Realistic biodiversity scenarios were developed based on long-term field surveys, and experimental assemblages replicated sequential loss of rare species which occurred across all trophic levels of these complex food webs. Response variables represented multiple components of ecosystem functioning, including nutrient cycling, primary and secondary production, organic matter accumulation and whole ecosystem metabolism. Species richness significantly affected ecosystem function, even after statistically controlling for potentially confounding factors such as total biomass and direct trophic interactions. Overall, loss of rare species was generally associated with lower nutrient concentrations, phytoplankton and zooplankton densities, and whole ecosystem metabolism when compared with more diverse assemblages. This pattern was also observed for overall ecosystem multifunctionality, a combined metric representing the ability of an ecosystem to simultaneously maintain multiple functions. One key exception was attributed to time-dependent effects of intraguild predation, which initially increased values for most ecosystem response variables, but resulted in decreases over time likely due to reduced nutrient remineralization by surviving predators. At the same time, loss of species did not result in strong trophic cascades, possibly a result of compensation and complexity of these multitrophic ecosystems along with a dominance of bottom-up effects. Our results indicate that although rare species may comprise minor components of communities, their loss can have profound ecosystem consequences across multiple trophic levels due to a combination of direct and indirect effects in diverse multitrophic ecosystems.     

Detritus, trophic dynamics and biodiversity

Traditional approaches to the study of food webs emphasize the transfer of local primary productivity in the form of living plant organic matter across trophic levels. However, dead organic matter, or detritus, a common feature of most ecosystems plays a frequently overlooked role as a dynamic heterogeneous resource and habitat for many species. We develop an integrative framework for understanding the impact of detritus that emphasizes the ontogeny and heterogeneity of detritus and the various ways that explicit inclusion of detrital dynamics alters generalizations about the structure and functioning of food webs. Through its influences on food web composition and dynamics, detritus often increases system stability and persistence, having substantial effects on trophic structure and biodiversity. Inclusion of detrital heterogeneity in models of food web dynamics is an essential new direction for ecological research.     

Intraguild predation promotes complex alternative states along a productivity gradient

Intraguild predation is the simplest, ubiquitous form of trophic omnivory, known to greatly influence the structure and functioning of natural and managed food webs. Although alternative states are fundamental to intraguild predation dynamics, only necessary conditions for alternative states have been previously reported. Using simple models, we found complex but systematic patterns in which different alternative states occur along a productivity gradient, and clarified the sufficient conditions to separate these patterns. We found that two quantities known to control the necessary conditions also determine the sufficient conditions: (1) relative energy transfer efficiency through alternative trophic pathways to an intraguild predator, and (2) relative resource exploitation ability between intraguild prey and predator. These governing quantities suggest how body size and stoichiometric relations between intraguild prey and predators can influence the possibility of alternative states. Our results indicate that food webs involving intraguild predation have a high potential of complex alternative states, and their management can be highly precarious.     

全球变化对食物网结构影响机制的研究进展

食物网主要依靠基于不同营养级间物种互作形成的上行与下行调控维持其结构。全球变化能够改变种间关系, 威胁生物多样性的维持, 然而目前对全球变化改变食物网结构的机制仍处于探索阶段。近年来通过大时空格局与多营养级食物网研究, 发现全球变化的作用机制主要可归结为3种: 物候错配、关键种丧失与生物入侵。该文聚焦于这3种机制, 综述各种机制造成的食物网结构变化并探讨相关的进化与生态驱动因素。三种干扰机制均通过改变原有种间关系, 影响食物网调控, 改变食物网结构。不同的是, 物候错配造成的种间关系变化是由于不同物种的物候对全球变化产生非同步响应所致; 关键种丧失则使营养级间取食/捕食关系发生变化甚至缺失; 而入侵物种通过竞争排除同营养级物种改变种间关系。最后, 该文提出食物网结构变化的实质是物种是否能够适应快速变化的生态环境, 并据此展望未来研究方向。随着全球变化影响日益加剧, 急需继续深入探索导致全球变化下食物网结构改变的机制, 为制定合理的生物多样性保护与生态修复规划提供重要理论支撑。     

The importance of omnivory and predator regulation of prey in freshwater fish assemblages of North America

Synopsis Food webs for 15 freshwater ecosystems in North America are reconstructed, based upon fish feeding habits in these ecosystems and established trophic categorizations for aquatic invertebrates. My own research in Goose Creek, Virginia, as well as literature data, provided the necessary information. One property of theoretical interest for food webs, namely omnivory, is examined, and the results are related to various topics more typically studied by freshwater fish ecologists, including predator-prey regulation, optimal foraging theory, and tropho-dynamics. The data indicate that omnivory is often important in freshwater ecosystems, such that fish may not always control the abundance of their prey. It is emphasized that knowledge of omnivory patterns can provide important insights into community structure and function in freshwater ecosystems.