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.     

Testing the ecological consequences of evolutionary change using elements

Understanding the ecological consequences of evolutionary change is a central challenge in contemporary biology. We propose a framework based on the ~25 elements represented in biology, which can serve as a conduit for a general exploration of poorly understood evolution‐to‐ecology links. In this framework, known as ecological stoichiometry, the quantity of elements in the inorganic realm is a fundamental environment, while the flow of elements from the abiotic to the biotic realm is due to the action of genomes, with the unused elements excreted back into the inorganic realm affecting ecological processes at higher levels of organization. Ecological stoichiometry purposefully assumes distinct elemental composition of species, enabling powerful predictions about the ecological functions of species. However, this assumption results in a simplified view of the evolutionary mechanisms underlying diversification in the elemental composition of species. Recent research indicates substantial intraspecific variation in elemental composition and associated ecological functions such as nutrient excretion. We posit that attention to intraspecific variation in elemental composition will facilitate a synthesis of stoichiometric information in light of population genetics theory for a rigorous exploration of the ecological consequences of evolutionary change.     

Invasive aquarium fish transform ecosystem nutrient dynamics

Trade of ornamental aquatic species is a multi-billion dollar industry responsible for the introduction of myriad fishes into novel ecosystems. Although aquarium invaders have the potential to alter ecosystem function, regulation of the trade is minimal and little is known about the ecosystem-level consequences of invasion for all but a small number of aquarium species. Here, we demonstrate how ecological stoichiometry can be used as a framework to identify aquarium invaders with the potential to modify ecosystem processes. We show that explosive growth of an introduced population of stoichiometrically unique, phosphorus (P)-rich catfish in a river in southern Mexico significantly transformed stream nutrient dynamics by altering nutrient storage and remineralization rates. Notably, changes varied between elements; the P-rich fish acted as net sinks of P and net remineralizers of nitrogen. Results from this study suggest species-specific stoichiometry may be insightful for understanding how invasive species modify nutrient dynamics when their population densities and elemental composition differ substantially from native organisms. Risk analysis for potential aquarium imports should consider species traits such as body stoichiometry, which may increase the likelihood that an invasion will alter the structure and function of ecosystems.     

Ecological stoichiometry and the shape of resource-based tradeoffs

Fitness tradeoffs are expected when organisms must allocate a limited resource between two or more traits. Despite the fact that most resources are made up of multiple elements or components, tradeoff models have traditionally been implicitly or explicitly based on a single currency. Here I propose a model for a resource-based tradeoff between two traits that share two resource elements, and suggest ecological stoichiometry as a critical mechanism controlling the shape of tradeoffs. The tradeoff curve in the model has different shapes depending on the elemental stoichiometry of the traits and the available resource. I further describe a conceptual framework that combines a tradeoff curve with a fitness function to predict an optimal combination of traits in stoichiometrically different environments. The model suggests that even though the fitness function is fixed, an optimal combination of traits can vary depending on the stoichiometry of both the traits and the supplied resource.     

Ecological Stoichiometry and Multi-element Transfer in a Coastal Ecosystem

Energy (carbon) flows and element cycling are fundamental, interlinked principles explaining ecosystem processes. The element balance in components, interactions and processes in ecosystems (ecological stoichiometry; ES) has been used to study trophic dynamics and element cycling. This study extends ES beyond its usual limits of C, N, and P and examines the distribution and transfer of 48 elements in 16 components of a coastal ecosystem, using empirical and modeling approaches. Major differences in elemental composition were demonstrated between abiotic and biotic compartments and trophic levels due to differences in taxonomy and ecological function. Mass balance modeling for each element, based on carbon fluxes and element:C ratios, was satisfactory for 92.5% of all element–compartment combinations despite the complexity of the ecosystem model. Model imbalances could mostly be explained by ecological processes, such as increased element uptake during the spring algal bloom. Energy flows in ecosystems can thus realistically estimate element transfer in the environment, as modeled uptake is constrained by metabolic rates and elements available. The dataset also allowed us to examine one of the key concepts of ES, homeostasis, for more elements than is normally possible. The relative concentrations of elements in organisms compared to their resources did not provide support for the theory that autotrophs show weak homeostasis and showed that the strength of homeostasis by consumers depends on the type of element (for example, macroelement, trace element). Large-scale, multi-element ecosystem studies are essential to evaluate and advance the framework of ES and the importance of ecological processes.     

Eco-evolutionary consequences of habitat warming and fragmentation in communities

Eco-evolutionary dynamics can mediate species and community responses to habitat warming and fragmentation, two of the largest threats to biodiversity and ecosystems. The eco-evolutionary consequences of warming and fragmentation are typically studied independently, hindering our understanding of their simultaneous impacts. Here, we provide a new perspective rooted in trade-offs among traits for understanding their eco-evolutionary consequences. On the one hand, temperature influences traits related to metabolism, such as resource acquisition and activity levels. Such traits are also likely to have trade-offs with other energetically costly traits, like antipredator defences or dispersal. On the other hand, fragmentation can influence a variety of traits (e.g. dispersal) through its effects on the spatial environment experienced by individuals, as well as properties of populations, such as genetic structure. The combined effects of warming and fragmentation on communities should thus reflect their collective impact on traits of individuals and populations, as well as trade-offs at multiple trophic levels, leading to unexpected dynamics when effects are not additive and when evolutionary responses modulate them. Here, we provide a road map to navigate this complexity. First, we review single-species responses to warming and fragmentation. Second, we focus on consumer–resource interactions, considering how eco-evolutionary dynamics can arise in response to warming, fragmentation, and their interaction. Third, we illustrate our perspective with several example scenarios in which trait trade-offs could result in significant eco-evolutionary dynamics. Specifically, we consider the possible eco-evolutionary consequences of (i) evolution in thermal performance of a species involved in a consumer–resource interaction, (ii) ecological or evolutionary changes to encounter and attack rates of consumers, and (iii) changes to top consumer body size in tri-trophic food chains. In these scenarios, we present a number of novel, sometimes counter-intuitive, potential outcomes. Some of these expectations contrast with those solely based on ecological dynamics, for example, evolutionary responses in unexpected directions for resource species or unanticipated population declines in top consumers. Finally, we identify several unanswered questions about the conditions most likely to yield strong eco-evolutionary dynamics, how better to incorporate the role of trade-offs among traits, and the role of eco-evolutionary dynamics in governing responses to warming in fragmented communities.     

Within-population variation in the chemistry of life: the stoichiometry of sexual dimorphism in multiple dimensions

In sexual species, phenotypic divergence between males and females, or sexual dimorphism, is often the source of the most staggering examples of phenotypic variation in nature. Theory suggests that exaggerated sexual traits should drive sex-specific nutritional demands. Advances in spectrometry enable rapid quantification of the elements that make up individuals and traits, which can be used to assess patterns of intraspecific variation and the contribution of nutritionally-demanding sexual traits to these patterns. We measured dimorphism in the whole body stoichiometry of Hyalella amphipods and examined whether nutritional demands of exaggerated sexual traits differ from those of similar traits not under sexual selection. We found striking sexual dimorphism in multivariate whole body elemental composition (i.e., the ionome), including elements important for organismal growth and performance. In males, the exaggerated, sexually-selected claw-like appendage (posterior gnathopod) differed significantly in mass-specific stoichiometry from a similarly sized and serially homologous non-sexual trait (fifth pereopod), indicating that there are fundamental differences in the construction of sexual traits in relation to similar traits that are not under sexual selection. While sexually selected traits do differ from non-sexual traits in their ionomes, we found that possessing an exaggerated trait does not change organismal stoichiometry, indicating that trait exaggeration may not be directly driving ionomic sexual dimorphism. Finally, we found that larger traits are not comparatively larger resource sinks for any element, suggesting that the possession of larger traits is not a function of greater allocation of resources. Together, we discovered substantial sexual dimorphism at the lowest level of organization, chemical elements. Such information illuminates predictions about dimorphisms in foraging behavior, nutritional physiology, and sex-specific selection on the underlying loci. High throughput, multidimensional data on sexual divergence in stoichiometric composition is a powerful tool in understanding the evolutionary ecology of sexual dimorphisms.     

生态化学计量学：复杂生命系统奥秘的探索

 20世纪以来,生物科学的发展异军突起,成为发展最快的学科,不仅学科分类逐渐细化,而且研究领域也逐渐深入,然而,这种分化和深入也可能会掩盖生物的一些最普遍特征。地球上的生物是否具有统一的、更本质的特征？能否把不同生物学领域和不同层次的知识联系起来？随着对这些问题的不断探索,一门新兴的学科——生态化学计量学,在最近20年悄然兴起。生态化学计量学结合了生物学、化学和物理学等基本原理,是研究生物系统能量平衡和多重化学元素（主要是C、N、P）平衡的科学。这一研究领域使得生物学科不同层次（分子、细胞、有机体、种群、生态系统和全球尺度）的研究理论能够有机地统一起来。目前,生态化学计量学已经广泛应用于种群动态、生物体营养动态、微生物营养、寄主_病原关系、生物共生关系、消费者驱动的养分循环、限制性元素的判断、生态系统比较分析和森林演替与衰退及全球C、N、P生物地球化学循环等研究中,并取得了许多研究成果。该文概述了生态化学计量的概念、历史起源和基本理论,重点介绍了生态化学计量学理论在消费者驱动的养分循环、限制性养分元素判别以及全球C、N、P循环等方面的应用进展,并对生态化学计量学未来的研究方向进行了展望,期望引起国内同行的重视并有助于推动我国在此领域开展相关研究。     

Stoichiometric distribution models: ecological stoichiometry at the landscape extent

Human activities are altering the fundamental geography of biogeochemicals. Yet we lack an understanding of how the spatial patterns in organismal stoichiometry affect biogeochemical processes and the tools to predict the impacts of global changes on biogeochemical processes. In this contribution we develop stoichiometric distribution models (StDMs), which allow us to map spatial structure in resource elemental composition across a landscape and evaluate spatial responses of consumers. We parameterise StDMs for a consumer‐resource (moose‐white birch) system and demonstrate that we can develop predictive models of resource stoichiometry across a landscape and that such models could improve our predictions of consumer space use. With results from our study system application, we argue that explicit consideration of the spatial patterns in organismal elemental composition may uncover emergent individual, population, community and ecosystem properties that are not revealed at the local extents routinely used in ecological stoichiometry. We discuss perspectives for further developments and application of StDMs to advance three emerging frameworks for spatial ecosystem ecology in an era of global change; meta‐ecosystem theory, macroecological stoichiometry and remotely sensed biogeochemistry. Progress on these emerging frameworks will allow for the integration of ecological stoichiometry and individual space use and fitness.     

Extending eco-evolutionary theory with oligomorphic dynamics

Understanding the interplay between ecological processes and the evolutionary dynamics of quantitative traits in natural systems remains a major challenge. Two main theoretical frameworks are used to address this question, adaptive dynamics and quantitative genetics, both of which have strengths and limitations and are often used by distinct research communities to address different questions. In order to make progress, new theoretical developments are needed that integrate these approaches and strengthen the link to empirical data. Here, we discuss a novel theoretical framework that bridges the gap between quantitative genetics and adaptive dynamics approaches. ‘Oligomorphic dynamics’ can be used to analyse eco-evolutionary dynamics across different time scales and extends quantitative genetics theory to account for multimodal trait distributions, the dynamical nature of genetic variance, the potential for disruptive selection due to ecological feedbacks, and the non-normal or skewed trait distributions encountered in nature. Oligomorphic dynamics explicitly takes into account the effect of environmental feedback, such as frequency- and density-dependent selection, on the dynamics of multi-modal trait distributions and we argue it has the potential to facilitate a much tighter integration between eco-evolutionary theory and empirical data.     

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.     

Can metabolic traits explain animal community assembly and functioning?

All animals on Earth compete for free energy, which is acquired, assimilated, and ultimately allocated to growth and reproduction. Competition is strongest within communities of sympatric, ecologically similar animals of roughly equal size (i.e. horizontal communities), which are often the focus of traditional community ecology. The replacement of taxonomic identities with functional traits has improved our ability to decipher the ecological dynamics that govern the assembly and functioning of animal communities. Yet, the use of low-resolution and taxonomically idiosyncratic traits in animals may have hampered progress to date. An animal's metabolic rate (MR) determines the costs of basic organismal processes and activities, thus linking major aspects of the multifaceted constructs of ecological niches (where, when, and how energy is obtained) and ecological fitness (how much energy is accumulated and passed on to future generations). We review evidence from organismal physiology to large-scale analyses across the tree of life to propose that MR gives rise to a group of meaningful functional traits – resting metabolic rate (RMR), maximum metabolic rate (MMR), and aerobic scope (AS) – that may permit an improved quantification of the energetic basis of species coexistence and, ultimately, the assembly and functioning of animal communities. Specifically, metabolic traits integrate across a variety of typical trait proxies for energy acquisition and allocation in animals (e.g. body size, diet, mobility, life history, habitat use), to yield a smaller suite of continuous quantities that: (1) can be precisely measured for individuals in a standardized fashion; and (2) apply to all animals regardless of their body plan, habitat, or taxonomic affiliation. While integrating metabolic traits into animal community ecology is neither a panacea to disentangling the nuanced effects of biological differences on animal community structure and functioning, nor without challenges, a small number of studies across different taxa suggest that MR may serve as a useful proxy for the energetic basis of competition in animals. Thus, the application of MR traits for animal communities can lead to a more general understanding of community assembly and functioning, enhance our ability to trace eco-evolutionary dynamics from genotypes to phenotypes (and vice versa), and help predict the responses of animal communities to environmental change. While trait-based ecology has improved our knowledge of animal communities to date, a more explicit energetic lens via the integration of metabolic traits may further strengthen the existing framework.     

Linking genes to communities and ecosystems: Daphnia as an ecogenomic model

How do genetic variation and evolutionary change in critical species affect the composition and functioning of populations, communities and ecosystems? Illuminating the links in the causal chain from genes up to ecosystems is a particularly exciting prospect now that the feedbacks between ecological and evolutionary changes are known to be bidirectional. Yet to fully explore phenomena that span multiple levels of the biological hierarchy requires model organisms and systems that feature a comprehensive triad of strong ecological interactions in nature, experimental tractability in diverse contexts and accessibility to modern genomic tools. The water flea Daphnia satisfies these criteria, and genomic approaches capitalizing on the pivotal role Daphnia plays in the functioning of pelagic freshwater food webs will enable investigations of eco-evolutionary dynamics in unprecedented detail. Because its ecology is profoundly influenced by both genetic polymorphism and phenotypic plasticity, Daphnia represents a model system with tremendous potential for developing a mechanistic understanding of the relationship between traits at the genetic, organismal and population levels, and consequences for community and ecosystem dynamics. Here, we highlight the combination of traits and ecological interactions that make Daphnia a definitive model system, focusing on the additional power and capabilities enabled by recent molecular and genomic advances.     

生态化学计量学在水生态系统中的研究与应用

生态化学计量学主要是研究碳、氮、磷等元素在各种生态过程中平衡的一门科学,其核心问题是揭示生物体元素组成的差异对生态功能的影响。由于生态化学计量学研究可以把生态实体的各个层次在元素水平上统一起来,因此生态化学计量学已成为许多生态系统的新兴研究工具。目前,生态化学计量学的研究与应用已深入到生态学的各个层次(分子、细胞、个体、种群、群落、生态系统及区域等不同尺度)。该文围绕生态化学计量学的两个重要组成理论,并结合笔者近年来的研究,归纳总结了生态化学计量学在水生态系统中的研究与应用及未来研究重点,希望有助于推动我国生态化学计量学在水生态系统中的应用研究。     

Sex,competition and mimicry: an eco-evolutionary model reveals unexpected impacts of ecological interactions on the evolution of phenotypes in sympatry

Phenotypic evolution in sympatric species can be strongly impacted by species interactions, either mutualistic or antagonistic. Heterospecific reproductive behaviours between sympatric species have been shown to favour phenotypic divergence of traits used as sexual cues. Those traits may also be involved in local adaptation or in other types of species interactions and, as a result, undergo complex evolutions across sympatric species. Here we focus on mimicry and study how reproductive interference may impair phenotypic convergence between species with various levels of defence. We use a deterministic model assuming two sympatric species where individuals can display two different warning colour patterns. This eco-evolutionary model explores how ecological interactions shape phenotypic evolution within sympatric species. We investigate the effect of 1) the opposing density-dependent selections exerted on colour patterns by predation and reproductive behaviour and 2) the impact of relative species and phenotype abundances on the fitness costs faced by each individual depending on their species and phenotype. Our model shows that reproductive interference may limit the convergent effect of mimetic interactions and may promote phenotypic divergence between Müllerian mimics. The divergent and convergent evolution of traits also strongly depends on the relative species and phenotype abundances and levels of trophic competition, highlighting how the eco-evolutionary feedbacks between phenotypic evolution and species abundances may result in strikingly different evolutionary routes.     

Eco-evolutionary dynamics in herbivorous insect communities mediated by induced plant responses

It is increasingly recognized that the ecology of communities and evolution of species within communities are interdependent, and researchers have been paying attention to this rapidly emerging field of research, i.e., through studies on eco-evolutionary dynamics. Most of the studies on eco-evolutionary dynamics have been concerned with direct trophic interactions. However, community ecologists have shown that trait-mediated indirect effects play an important role in shaping the structure of natural communities. In particular, in terrestrial plant–insect systems, indirect effects mediated through herbivore-induced plant responses are common and have a great impact on the structure of herbivore communities. This review describes eco-evolutionary dynamics in herbivorous insect communities, and specifically focuses on the key role of herbivore-induced plant responses in eco-evolutionary dynamics. First, I review studies on the evolution of herbivore traits relevant to plant induction and discuss evolution in a community context mediated by induced plant responses. Second, I highlight how intraspecific genetic variation or evolution in herbivore traits can influence herbivore community structure. Finally, I propose the hypothetical model that induced plant responses supports eco-evolutionary feedback in herbivore communities. In this review, I argue that the application of the indirect interaction web approaches into studies on eco-evolutionary will provide profound insights into understanding of mechanisms of the generation and maintenance of biodiversity.     

Phenotypic Variability: Its Components, Measurement and Underlying Developmental Processes

Variability contrasts with variation in that variability describes the potential for variation, not simply the expressed variation. The power of studying variability lies in creating a conceptual framework around which the relationship between the genotype and phenotype can be understood. Here, we attempt to demonstrate the importance of phenotypic variability, how it structures variation, and how fundamental developmental processes structure variability. Given the broad scope of this topic, we focus on three widely studied properties of variability: canalization, developmental stability and morphological integration. We have organized the paper to emphasize the importance of differentiating between the theory surrounding these components of phenotypic variability, their measurement and the biological factors surrounding their expression. First, we define these properties of variability, how they relate to each other and to variability as a whole. Second, we summarize the common methods of measurement for canalization, developmental stability and morphological integration and the reasoning behind these methods. Finally, we focus on jaw development as an example of how the basic processes of development affect variability and the resultant variation, with emphasis on how processes at all levels of the organismal hierarchy interact with one another and contribute to phenotypic variability.     

Coevolution of slow-fast populations: evolutionary sliding, evolutionary pseudo-equilibria and complex Red Queen dynamics

We study the interplay of ecological and evolutionary dynamics in communities composed of populations with contrasting time-scales. In such communities, genetic variation of individual traits can cause population transitions between stationary and cyclic ecological regimes, hence abrupt variations in fitness. Such abrupt variations raise ridges in the adaptive landscape, where the populations are poised between equilibrium and cyclic coexistence and along which evolutionary trajectories can remain sliding for long times or halt at special points called evolutionary pseudo-equilibria. These novel phenomena should be generic to all systems in which ecological interactions cause fitness to vary discontinuously. They are demonstrated by the analysis of a predator-prey community, with one adaptive trait for each population. The eco-evolutionary dynamics of the system show a number of other distinctive features, including evolutionary extinction and two forms of Red Queen dynamics. One of them is characterized by intermittent bouts of cyclic oscillations of the two populations.