Rapid evolution of a consumer stoichiometric trait destabilizes consumer–producer dynamics

Recent studies have shown that adaptive evolution can be rapid enough to affect contemporary ecological dynamics in nature (i.e. ‘rapid evolution’). These studies tend to focus on trait functions relating to interspecific interactions; however, the importance of rapid evolution of stoichiometric traits has been relatively overlooked. Various traits can affect the balance of elements (carbon, nitrogen, and phosphorus) of organisms, and rapid evolution of such stoichiometric traits will not only alter population and community dynamics but also influence ecosystem functions such as nutrient cycling. Multiple environmental changes may exert a selection pressure leading to adaptation of stoichiometrically important traits, such as an organism's growth rate. In this paper, we use theoretical approaches to explore the connections between rapid evolution and ecological stoichiometry at both the population and ecosystem level. First, we incorporate rapid evolution into an ecological stoichiometry model to investigate the effects of rapid evolution of a consumer's stoichiometric phosphorus:carbon ratio on consumer–producer population dynamics. We took two complementary approaches, an asexual clonal genotype model and a quantitative genetic model. Next, we extended these models to explicitly track nutrients in order to evaluate the effect of rapid evolution at the ecosystem level. Our model results indicate rapid evolution of the consumer stoichiometric trait can cause complex dynamics where rapid evolution destabilizes population dynamics and rescues the consumer population from extinction (evolutionary rescue). The model results also show that rapid evolution may influence the level of nutrients available in the environment and the flux of nutrients across trophic levels. Our study represents an important step for theoretical integration of rapid evolution and ecological stoichiometry.     

A General Method for Modeling Population Dynamics and Its Applications

Studying populations, be it a microbe colony or mankind, is important for understanding how complex systems evolve and exist. Such knowledge also often provides insights into evolution, history and different aspects of human life. By and large, populations’ prosperity and decline is about transformation of certain resources into quantity and other characteristics of populations through growth, replication, expansion and acquisition of resources. We introduce a general model of population change, applicable to different types of populations, which interconnects numerous factors influencing population dynamics, such as nutrient influx and nutrient consumption, reproduction period, reproduction rate, etc. It is also possible to take into account specific growth features of individual organisms. We considered two recently discovered distinct growth scenarios: first, when organisms do not change their grown mass regardless of nutrients availability, and the second when organisms can reduce their grown mass by several times in a nutritionally poor environment. We found that nutrient supply and reproduction period are two major factors influencing the shape of population growth curves. There is also a difference in population dynamics between these two groups. Organisms belonging to the second group are significantly more adaptive to reduction of nutrients and far more resistant to extinction. Also, such organisms have substantially more frequent and lesser in amplitude fluctuations of population quantity for the same periodic nutrient supply (compared to the first group). Proposed model allows adequately describing virtually any possible growth scenario, including complex ones with periodic and irregular nutrient supply and other changing parameters, which present approaches cannot do.     

Vectors with autonomy: what distinguishes animal‐mediated nutrient transport from abiotic vectors?

Animal movements are important drivers of nutrient redistribution that can affect primary productivity and biodiversity across various spatial scales. Recent work indicates that incorporating these movements into ecosystem models can enhance our ability to predict the spatio‐temporal distribution of nutrients. However, the role of animal behaviour in animal‐mediated nutrient transport (i.e. active subsidies) remains under‐explored. Here we review the current literature on active subsidies to show how the behaviour of active subsidy agents makes them both ecologically important and qualitatively distinct from abiotic processes (i.e. passive subsidies). We first propose that animal movement patterns can create similar ecological effects (i.e. press and pulse disturbances) in recipient ecosystems, which can be equal in magnitude to or greater than those of passive subsidies. We then highlight three key behavioural features distinguishing active subsidies. First, organisms can transport nutrients counter‐directionally to abiotic forces and potential energy gradients (e.g. upstream). Second, unlike passive subsidies, organisms respond to the patterns of nutrients that they generate. Third, animal agents interact with each other. The latter two features can form positive‐ or negative‐feedback loops, creating patterns in space or time that can reinforce nutrient hotspots in places of mass aggregations and/or create lasting impacts within ecosystems. Because human‐driven changes can affect both the space‐use of active subsidy species and their composition at both population (i.e. individual variation) and community levels (i.e. species interactions), predicting patterns in nutrient flows under future modified environmental conditions depends on understanding the behavioural mechanisms that underlie active subsidies and variation among agents' contributions. We conclude by advocating for the integration of animal behaviour, animal movement data, and individual variation into future conservation efforts in order to provide more accurate and realistic assessments of changing ecosystem function.     

Daisyworld is Darwinian: constraints on adaptation are important for planetary self-regulation

The Daisyworld model demonstrates that self-regulation of the global environment can emerge from competition amongst types of life altering their local environment in different ways. Robertson & Robinson (1998. J. theor. Biol.195, 129-134) presented what they describe as a "Darwinian Daisyworld" in which the ability of organisms to adapt their internal physiology in response to environmental change undermines their ability to regulate their environment. They assume that there are no bounds on the environmental conditions that organisms can adapt to and that equal growth rates can potentially be achieved under any conditions. If adaptation could respond sufficiently rapidly to changes in the environment, this would eliminate any need for the environment to be regulated in the first place, because all possible states of the environment would be equally tolerable to life. However, the thermodynamics, chemistry and structure of living organisms set bounds on the range of environmental conditions that can be adapted to. As these bounds are approached, environmental conditions limit growth rate, and adaptations necessary for survival can also cost energy. Here we take account of such constraints and find that environmental regulation is recovered in the Daisyworld model. Hence, we suggest that constraints are an important part of a self-regulating planetary system.     

Environmental fluctuations promote intraspecific diversity and population persistence via inflationary effects

The impact of temporal variation in the environment, specifically the amount of temporal autocorrelation, on population processes is of growing interest in ecology and evolutionary biology. It was recently discovered that temporal autocorrelation in the environment can significantly increase the abundance of populations that would otherwise have low, or even negative long‐term growth rates (via so‐called ‘inflationary effects’), provided that immigration from another source prevents extinction. Here we use a mathematical model to ask whether inflationary effects can also increase population persistence without immigration if different phenotypes within that population partition growth over time and buffer each other from extinction via mutation. Using a combination of analytical and numerical methods, we find that environmental autocorrelation can inflate the abundance of phenotypes that would otherwise be excluded from the population, provided that phenotypes are sufficiently different in their use of the environment. This inflation of abundance at the phenotypic level also generates an inflation of abundance at the population level. Remarkably, intraspecific inflationary effects can increase both phenotypic and whole population abundance even if one or all phenotypes are maladapted to the environment, as long as mutations prevent phenotypic extinction during periods of poor environmental conditions. Given the prevalence of temporally autocorrelated environmental variables in nature, intraspecific inflationary effects have the potential to be of widespread importance for population persistence as well as the maintenance of intraspecific diversity.     

Impacts of parasitic plants on natural communities

Parasitic plants have profound effects on the ecosystems in which they occur. They are represented by some 4000 species and can be found in most major biomes. They acquire some or all of their water, carbon and nutrients via the vascular tissue of the host's roots or shoots. Parasitism has major impacts on host growth, allometry and reproduction, which lead to changes in competitive balances between host and nonhost species and therefore affect community structure, vegetation zonation and population dynamics. Impacts on hosts may further affect herbivores, pollinators and seed vectors, and the behaviour and diversity of these is often closely linked to the presence and abundance of parasitic plants. Parasitic plants can therefore be considered as keystone species. Community impacts are mediated by the host range of the parasite (the diversity of species that can potentially act as hosts) and by their preference and selection of particular host species. Parasitic plants can also alter the physical environment around them--including soil water and nutrients, atmospheric CO2 and temperature--and so may also be considered as ecosystem engineers. Such impacts can have further consequences in altering the resource supply to and behaviour of other organisms within parasitic plant communities.     

A rule-based approach to the modelling of bacterial ecosystems

This paper presents an approach to ecological/evolutionary modelling that is inspired by natural bacterial ecosystems and bacterial evolution. An individual-based artificial ecosystem model is proposed, which is designed to explore the evolvability of adaptive behavioural strategies in artificial bacteria represented by rule-based learning classifier systems. The proposed ecosystem model consists of a n-dimensional environmental grid, which can contain different types of artificial resources in arbitrary arrangements. The resources provide the energy that is necessary for the organisms to sustain life, and can trigger different types of behaviour in the organisms, such as movement towards nutrients and away from toxic substances, growth, and the controlled release of signalling resources. The balance between energy and material is modelled carefully to ensure that the ecosystem is dissipative. Those organisms that are able to efficiently exploit the available resources gradually accumulate enough energy to reproduce (by binary fission) and generate copies of themselves in the environment. Organisms are also able to produce their own resources, which can potentially be used as markers to send signals to other organisms (a behaviour known as quorum sensing). The complex relationships between stimuli and actions in the organisms are stochastically altered by means of mutations, thus enabling the organisms to adapt to their environment and maximise their lifespan and reproductive success. In this paper, the proposed bacterial ecosystem model is defined formally and its structure is discussed in detail. This is followed by results from simulation experiments that illustrate the model's operation and how it can be used in evolutionary modelling/computing scenarios.     

Evolutionary dynamics and highly optimized tolerance

We develop a numerical model of a lattice community based on Highly Optimized Tolerance (HOT), which relates the evolution of complexity to robustness tradeoffs in an uncertain environment. With the model, we explore scenarios for evolution and extinction which are abstractions of processes which are commonly discussed in biological and ecological case studies. These include the effects of different habitats on the phenotypic traits of the organisms, the effects of different mutation rates on adaptation, fitness, and diversity, and competition between generalists and specialists. The model exhibits a wide variety of microevolutionary and macroevolutionary phenomena which can arise in organisms which are subject to random mutation, and selection based on fitness evaluated in a specific environment. Generalists arise in uniform habitats, where different disturbances occur with equal frequency, while specialists arise when the relative frequency of different disturbances is skewed. Fast mutators are seen to play a primary role in adaptation, while slow mutators preserve well-adapted configurations. When uniform and skewed habitats are coupled through migration of the organisms, we observe a primitive form of punctuated equilibrium. Rare events in the skewed habitat lead to extinction of the specialists, whereupon generalists invade from the uniform habitat, adapt to their new surroundings, ultimately leading their progeny to become vulnerable to extinction in a subsequent rare disturbance.     

Evolutionary regime shifts in simulated ecosystems

Catastrophic regime shifts in ecosystems occur when the system is tipped into a new attractor state under some external forcing. Here we consider whether evolutionary adaptations within ecosystems can trigger similar transitions. We use an individual‐based, evolutionary model of interconnected ecosystems to analyze nonlinear changes in global state resulting from local adaptations. Transitions between periods of stability occur when new traits arise that allow exploitation of under‐utilized resources. Subsequent rapid growth of the population carrying the new trait causes abrupt environmental change that drives incumbent species extinct. We call these transitions ‘evolutionary regime shifts’. These internally generated perturbations can result in ecosystem collapse, followed by recovery to an alternate stable state, or occasionally system‐wide extinction. While these disruptions may have a negative impact on ecosystem productivity in individual simulation runs, mean results over many simulations show a trend for increasing ecosystem productivity and stability over time. Feedback between life and the abiotic environment in the model creates a ‘long‐tailed’ distribution of extinction sizes without any external trigger for large extinction events.     

Evolution of local facilitation in arid ecosystems

In harsh environments, sessile organisms can make their habitat more hospitable by buffering environmental stress or increasing resource availability. Although the ecological significance of such local facilitation is widely established, the evolutionary aspects have been seldom investigated. Yet addressing the evolutionary aspects of local facilitation is important because theoretical studies show that systems with such positive interactions can exhibit alternative stable states and that such systems may suddenly become extinct when they evolve (evolutionary suicide). Arid ecosystems currently experience strong changes in climate and human pressures, but little is known about the effects of these changes on the selective pressures exerted on the vegetation. Here, we focus on the evolution of local facilitation in arid ecosystems, using a lattice-structured model explicitly considering local interactions among plants. We found that the evolution of local facilitation depends on the seed dispersal strategy. In systems characterized by short-distance seed dispersal, adaptation to a more stressful environment leads to high local facilitation, allowing the population to escape extinction. In contrast, systems characterized by long-distance seed dispersal become extinct under increased stress even when allowed to adapt. In this case, adaptation in response to climate change and human pressures could give the final push to the desertification of arid ecosystems.     

Diel patterns of leaf and root growth: endogenous rhythmicity or environmental response?

Plants are sessile organisms forced to adjust to their surrounding environment. In a single plant the photoautotrophic shoot is exposed to pronounced environmental variations recurring in a day-night 24?h (diel) cycle, whereas the heterotrophic root grows in a temporally less fluctuating environment. The contrasting habitats of shoots and roots are reflected in different diel growth patterns and their responsiveness to environmental stimuli. Differences between diel leaf growth patterns of mono- and dicotyledonous plants correspond to their different organization and placement of growth zones. In monocots, heterotrophic growth zones are organized linearly and protected from the environment by sheaths of older leaves. In contrast, photosynthetically active growth zones of dicot leaves are exposed directly to the environment and show characteristic, species-specific diel growth patterns. It is hypothesized that the different exposure to environmental constraints and simultaneously the sink/source status of the growing organs may have induced distinct endogenous control of diel growth patterns in roots and leaves of monocot and dicot plants. Confronted by strong temporal fluctuations in environment, the circadian clock may facilitate robust intrinsic control of leaf growth in dicot plants.     

Plant diversity and insect herbivores: effects of environmental change in contrasting model systems

There is increasing concern over the potential impact of anthropogenic factors (e.g. increasing nutrient inputs, global climate change) on the rate of loss of diversity in ecosystems. Such losses may affect ecosystem processes. In addition, a change in diversity of one group of organisms may influence the diversity of species of the next trophic level. We examined the extent to which plant species richness influences that of insect herbivores in two systems: a long‐term field experiment on heather moorland and a model community in the Ecotron controlled environment facility. We examined the response of these two plant communities to environmental change, specifically increased levels of nutrients, grazing and atmospheric CO₂. We measured the indirect effects of changes in these factors on insect herbivores, both above‐ and below‐ground. In the moorland system, grazing was the largest influence on plant community structure. The community was dominated by one species, Calluna vulgaris, and loss of cover under heavy grazing allowed competing species to invade. However, grazing regime was not a major influence on the species richness of the insect herbivore community. Site was more important: there were a greater number of Hemiptera species on sites with more mineral soils than on peat sites, possibly because a greater variety of grass and herb species was present on the former sites. In the Ecotron, below‐ground factors were also important drivers of community change: elevated CO₂ increased carbon availability in the soil and there were simultaneous changes in the community composition of soil biota. Above‐ground, some plant species increased in abundance and others decreased, leading to interaction‐specific effects on the insect herbivores. In two very different studies of the effects of environmental change on the interactions between plants and their herbivores, several similar conclusions can be drawn: (1) effects are likely to be site‐ and interaction‐specific; (2) outcomes are likely to be strongly dependent on the initial state and the dominant species of the plant community; and (3) indirect effects, often mediated by below‐ground factors, may have a bigger influence on insect‐plant interactions than more direct effects of above‐ground factors.     

Regime shift and robustness of organism-created environments: A model for microbial ecosystems

Organism-environment interactions are different from organism-resource interactions in two respects: (1) resources can only be consumed by organisms whereas environmental conditions can be increased or decreased depending on the species; (2) high resource conditions generally stimulate the growth of organisms, whereas extreme environmental conditions are not necessarily favored because each species usually has an optimum range for growth. To investigate the properties of an organism-environment feedback system, we analyze a model for microbial ecosystems in which a single microorganism species can modify the environmental pH. We demonstrate that the equilibrium level of the environmental pH can be partially regulated at a relatively constant value even if the pH in the influx to the ecosystem changes over a wide range. For species that acidify the medium, the equilibrium pH is somewhat lower than the pH optimal for the species. The pH-stabilizing effect of microorganisms is stronger if their growth is self-limited by the environmental pH. When the influx becomes sufficiently alkaline, the population of the organism suddenly disappears and the environmental pH changes abruptly. The system shows bi-stability and hysteresis and therefore differs from a standard resource competition model composed of a single species that consumes resources.     

植物寄生对生态系统结构和功能的影响

寄生植物是生态系统中的特殊类群之一。植物寄生可以驱动生态系统中生物与非生物因子的变化,在生态系统结构与功能中起关键作用。寄生植物可以通过对寄主营养的集聚、改变凋落物的质量与数量、改变根的周转与分泌物格局、改变土壤水势,从而影响土壤理化特性。寄生植物会改变寄主的行为,改变寄主与非寄主植物之间的相互作用,从而影响植物群落的结构、多样性和动态,进而影响植被演替和植被生产力等。寄生植物与寄主均可被消费者取食,可直接或间接地影响生态系统的食草动物,包括草食昆虫等。寄生植物与寄主的其它寄生物存在竞争关系,可以直接或间接地影响寄主的其它寄生植物或病原真菌。寄生植物可以明显地改变土壤地球化学循环,将固有的不可动的成分转变为可利用的营养成分,改变土壤生物群落的结构与功能,从而显著影响地下生物群落。这些表明,植物寄生对生态系统的结构和功能有重要影响。针对特殊的被入侵的植物群落,该地寄生植物可以通过影响入侵植物寄主的生长、繁殖、生物量分配格局,改变土壤的理化特性,促进非寄主的非优势本地植物的生长,从而改变被入侵植物群落结构与多样性,达到生物防治及生态恢复的目的。     

微塑料污染对浮游生物影响研究进展

微塑料作为一种新型的环境污染物,大量存在于水环境中,给水生生物带来了极大的危害。浮游生物是水生食物链的基础,是水生生态系统物质循环和能量流动的重要环节;同时,浮游生物也是对各种环境污染物最敏感的类群。了解微塑料对浮游生物的影响是评价其生态风险的重要依据。本文介绍了环境中微塑料来源、特征及水生态系统微塑料污染现状,阐述了微塑料对水生生物的直接和间接危害,并重点聚焦于浮游植物和浮游动物,从个体、种群和群落的层次详细总结了微塑料的影响及其作用机制。最后,本文指出当前针对浮游生物微观基因和蛋白质组学,以及宏观种群和群落响应等方面的研究还非常缺乏,为今后开展微塑料危害研究提供参考。     

Non-linear biological responses to environmental noise affect population extinction risk

Non-linearities are commonly observed in the responses of organisms to environment. They potentially modify the qualitative and quantitative properties of population dynamics. We studied how non-linear responses to environment, or "noise filters", influence population variability and extinction risk by introducing coloured noise to the growth rate in the Hassell single-species model. The consequences of noise filtering were analysed by comparing the model dynamics with linearly filtered and non-linearly filtered noise that have the same mean. Three biologically plausible filters we used: saturating, unimodal optimum type, and sigmoid responses.
Filtering can either decrease or increase population variability when compared to linear noise response. The effect of noise filtering on variability is most pronounced with stable population dynamics and the outcome depends on the filter type, population growth rate, and noise colour.
Non-linear noise filtering predominantly increases extinction risks when population growth rate is low (R<5). As an exception, saturating filter has a window of decreased risk at very low growth rate and reddened environment. In the unstable range of the dynamics (15These results suggest that accounting for the non-linear responses to environment should be considered when estimating extinction risks and population variability. Moreover, the non-linear responses make noise colour a more important factor in these analyses.     

Pyrodiversity promotes interaction complementarity and population resistance

Theory predicts that network characteristics may help anticipate how populations and communities respond to extreme climatic events, but local environmental context may also influence responses to extreme events. For example, altered fire regimes in many ecosystems may significantly affect the context for how species and communities respond to changing climate. In this study, I tested whether the responses of a pollinator community to extreme drought were influenced by the surrounding diversity of fire histories (pyrodiversity) which can influence their interaction networks via changing partner availability. I found that at the community level, pyrodiverse landscapes promote functional complementarity and generalization, but did not consistently enhance functional redundancy or resistance to simulated co‐extinction cascades. Pyrodiversity instead supported flexible behaviors that enable populations to resist perturbations. Specifically, pollinators that can shift partners and network niches are better able to take advantage of the heterogeneity generated by pyrodiversity, thereby buffering pollinator populations against changes in plant abundances. These findings suggest that pyrodiversity is unlikely to improve community‐level resistance to droughts, but instead promotes population resistance and community functionality. This study provides unique evidence that resistance to extreme climatic events depends on both network properties and historical environmental context.     

Community extinction patterns in coloured environments

Understanding community responses to environmental variation is a fundamental aspect of ecological research, with direct ecological, conservation and economic implications. Here, we examined the role of the magnitude, correlation and autocorrelation structures of environmental variation on species' extinction risk (ER), and the probability of actual extinction events in model competitive communities. Both ER and probability increased with increasing positive autocorrelation when species responded independently to the environment, yet both decreased with a strong correlation between species-specific responses. These results are framed in terms of the synchrony between--and magnitude of variation within--species population sizes and are explained in terms of differences in noise amplification under different conditions. The simulation results are robust to changes in the strength of interspecific density dependence, and whether noise affects density-independent or density-dependent population processes. Similar patterns arose under different ranges of noise severity when these different model assumptions were examined. We compared our results with those from an analytically derived solution, which failed to capture many features of the simulation results.     

On the importance of dimensionality of space in models of space-mediated population persistence

Spatially explicit models have become widely used in today's mathematical ecology to study persistence of populations. For the sake of simplicity, population dynamics is often analyzed with 1-D models. An important question is: how adequate is such 1-D simplification of 2-D (or 3-D) dynamics for predicting species persistence. Here we show that dimensionality of the environment can play a critical role in the persistence of predator-prey interactions. We consider 1-D and 2-D dynamics of a predator-prey model with the prey growth damped by the Allee effect. We show that adding a second space coordinate into the 1-D model results in a pronounced increase of size of the domain in the parametric space where predator-prey coexistence becomes possible. This result is due to the possibility of formation of a number of 2-D patterns, which is impossible in the 1-D model. The 1-D and the 2-D models exhibit different qualitative responses to variations of system parameters. We show that in ecosystems having a narrow width (e.g. mountain valleys, vegetation patterns along canals in dry areas, etc.), extinction of species is more probable compared to ecosystems having a pronounced second dimension. In particular, the width of a long narrow natural reserve should be large enough to guarantee nonextinction of species via interaction of 2-D population patches.     

Recent advances in ecological stoichiometry: insights for population and community ecology

Conventional theories of population and community dynamics are based on a single currency such as number of individuals, biomass, carbon or energy. However, organisms are constructed of multiple elements and often require them (in particular carbon, phosphorus and nitrogen) in different ratios than provided by their resources; this mismatch may constrain the net transfer of energy and elements through trophic levels. Ecological stoichiometry, the study of the balance of elements in ecological processes, offers a framework for exploring ecological effects of such constraints. We review recent theoretical and empirical studies that have considered how stoichiometry may affect population and community dynamics. These studies show that stoichiometric constraints can affect several properties of populations (e.g. stability, oscillations, consumer extinction) and communities (e.g. coexistence of competitors, competitive interactions between different guilds). We highlight gaps in general knowledge and focus on areas of population and community ecology where incorporation of stoichiometric constraints may be particularly fruitful, such as studies of demographic bottlenecks, spatial processes, and multi-species interactions. Finally, we suggest promising directions for new research by recommending potential study systems (terrestrial insects, detritivory-based webs, soil communities) to improve our understanding of populations and communities. Our conclusion is that a better integration of stoichiometric principles and other theoretical approaches in ecology may allow for a richer understanding of both population and community structure and dynamics.