Spatial metrics as indicators of biodegradation benefits from bacterial dispersal networks

Ecological dynamics often show intricate variations in response to different spatial configurations of environmental conditions. For instance, efficient turnover of natural or anthropogenic compounds in soils strongly depends on the bioavailability of these compounds to metabolically active bacteria. Experimental and modelling studies have highlighted that fungal networks may considerably enhance bioavailability by facilitating bacterial dispersal. Therefore, such dispersal networks may play a key role in many soil processes, for example, in contaminant degradation. Particularly, simulation studies revealed that the spatial configurations of networks may be a crucial factor determining the bacterial access to contaminants. Since these spatial configurations are typically complex and not precisely known, suitable metrics describing them in an aggregated manner are required for assessing expected biodegradation benefits from different bacterial dispersal networks. Using a spatially explicit microbial model we randomly created various dispersal network configurations and simulated the resulting bacterial degradation of organic compounds. We investigated numerous spatial metrics for characterizing the manifold network configurations, and identified appropriate metrics based on nonparametric correlation measures. Our results show that single metrics can approximately indicate biodegradation performance, and that well-chosen combinations of two metrics offer very good assessments. Thus, our analysis provides characteristics to focus on when dealing with real fungal networks in future practical applications in environmental management. Moreover, the protocol we employed for deriving the appropriate metrics is suited to be adapted to other ecological studies of functional responses to spatial environmental characteristics, for instance, changes in ecosystem services or biodiversity aspects due to habitat loss and fragmentation.     

Variable Effects of Dispersal on Productivity of Bacterial Communities Due to Changes in Functional Trait Composition

Previous studies have shown variable relationships between dispersal rate and ecosystem functioning, but the reasons for and mechanisms behind variable dispersal rate – functioning patterns are currently unknown. In this study we used six bacterial lake water communities in a laboratory experiment in order to investigate how dispersal among communities influences community productivity by evaluating three different mechanisms: 1) changes in taxonomic diversity, 2) changes in phylogenetic diversity or 3) changes in the composition of functional traits. The experiment was conducted in two phases; (A) a dialysis bag experiment where the dispersal rate among six communities was manipulated and the subsequent change in bacterial diversity and growth rate was recorded, and (B) a regrowth experiment where we manipulated available resources to study how well a taxon grows on certain organic carbon resources, i.e. their functional traits. From experiment (B) we could thus estimate changes in functional traits in communities in experiment (A). Bacterial production was affected by dispersal, but not consistently among lakes. Neither change in taxonomic or phylogenetic diversity with dispersal could explain the observed dispersal – productivity relationships. Instead, changes in trait composition with dispersal, especially the communities’ ability to use p-coumaric acid, an aromatic compound, could explain the observed dispersal – productivity relationships. Changes in this trait caused by dispersal seemed especially important for bacterial productivity in waters with a high aromaticity of the organic matter pool. We conclude that the effect of dispersal on bacterial communities can affect ecosystem functioning in different ways, through changes in functional key-traits which are important for the local environment.     

面向多重生态保护目标的广东省生态系统服务退化风险情景模拟

稳定、协调、可持续的生态系统服务供给是维持区域自然生命系统、保障区域生态安全的基础。面临生态系统所受的人为与自然干扰,有必要准确评估生态系统服务退化风险,揭示区域生态安全和可持续性提升路径。现有研究多叠加所有生态系统服务以评估生态系统服务退化风险,未能面向区域生态问题或生态需求对不同类型生态系统服务进行整合。研究提出面向多重生态保护目标的生态系统服务退化风险情景模拟框架,考虑广东省生物多样性保育、水资源安全、自然灾害防范等生态保护目标,选取代表性生态系统服务类型(生境维持、水源涵养、水质净化、土壤保持、洪水缓解、热带气旋缓解),基于不同典型浓度路径情景评估2018—2035年生态系统服务退化风险。结果表明,2018—2035年广东省生态系统服务供给空间格局基本稳定,尽管有所退化、空间异质性强。生态系统服务退化风险的情景模拟表明,面向生物多样性保育目标的生态系统服务退化高风险区位于珠三角的深圳市、佛山市和江门市,以及粤西和粤东地区的湛江市、茂名市和揭阳市;面向水资源安全目标的生态系统服务退化高风险区位于珠三角的深圳市、中山市和江门市,以及粤西与粤东地区的茂名市、揭阳市和江门市;面向自然灾...     

Dispersal mood revealed by shifts from routine to direct flights in the meadow brown butterfly Maniola jurtina

A comprehensive mechanistic approach to dispersal requires the translation of the whole mobility register of the target organism into movement rules that could subsequently be used to model its displacements. According to the optimality paradigm, this procedure implies a cost–benefit analysis of mobility patterns taking into account not only movements, but also their external context and the internal state of the moving individuals. Using this framework, we detected a ‘dispersal mood’ in some individuals of the meadow brown butterfly Maniola jurtina. These adopted a direct flight strategy, which was topologically different from the previously documented foray search strategy. Those individuals that used the direct flight strategy moved straighter as soon as they left the habitat and avoided heading back to their patch of origin, which is the best inter‐patch search strategy when dispersal risks and costs are high. The direct flight strategy was conditional to sex: females used it twice as much as males. We suggest that this sex bias was due to female investment in offspring, which is maximized by male avoidance and spatial bet hedging. Inter‐patch dispersal of gravid females is crucial for the persistence of M. jurtina populations in spatially and temporally unpredictable environments.     

Evolutionary stability of ideal free dispersal strategies in patchy environments

A central question in the study of the evolution of dispersal is what kind of dispersal strategies are evolutionarily stable. Hastings (Theor Pop Biol 24:244-251, 1983) showed that among unconditional dispersal strategies in a spatially heterogeneous but temporally constant environment, the dispersal strategy with no movement is convergent stable. McPeek and Holt's (Am Nat 140:1010-1027, 1992) work suggested that among conditional dispersal strategies in a spatially heterogeneous but temporally constant environment, an ideal free dispersal strategy, which results in the ideal free distribution for a single species at equilibrium, is evolutionarily stable. We use continuous-time and discrete-space models to determine when the dispersal strategy with no movement is evolutionarily stable and when an ideal free dispersal strategy is evolutionarily stable, both in a spatially heterogeneous but temporally constant environment.     

Evolutionarily Stable and Convergent Stable Strategies in Reaction–Diffusion Models for Conditional Dispersal

We consider a mathematical model of two competing species for the evolution of conditional dispersal in a spatially varying, but temporally constant environment. Two species are different only in their dispersal strategies, which are a combination of random dispersal and biased movement upward along the resource gradient. In the absence of biased movement or advection, Hastings showed that the mutant can invade when rare if and only if it has smaller random dispersal rate than the resident. When there is a small amount of biased movement or advection, we show that there is a positive random dispersal rate that is both locally evolutionarily stable and convergent stable. Our analysis of the model suggests that a balanced combination of random and biased movement might be a better habitat selection strategy for populations.     

On several conjectures from evolution of dispersal

We address several conjectures raised in Cantrell et al. [Evolution of dispersal and ideal free distribution, Math. Biosci. Eng. 7 (2010), pp. 17-36 [ 9 ]] concerning the dynamics of a diffusion-advection-competition model for two competing species. A conditional dispersal strategy, which results in the ideal free distribution of a single population at equilibrium, was found in Cantrell et al. [ 9 ]. It was shown in [ 9 ] that this special dispersal strategy is a local evolutionarily stable strategy (ESS) when the random diffusion rates of the two species are equal, and here we show that it is a global ESS for arbitrary random diffusion rates. The conditions in [ 9 ] for the coexistence of two species are substantially improved. Finally, we show that this special dispersal strategy is not globally convergent stable for certain resource functions, in contrast with the result from [ 9 ], which roughly says that this dispersal strategy is globally convergent stable for any monotone resource function.     

Regional zooplankton dispersal provides spatial insurance for ecosystem function

Changing environmental conditions are affecting diversity and ecosystem function globally. Theory suggests that dispersal from a regional species pool may buffer against changes in local community diversity and ecosystem function after a disturbance through the establishment of functionally redundant tolerant species. The spatial insurance provided by dispersal may decrease through time after environmental change as the local community monopolizes resources and reduces community invasibility. To test for evidence of the spatial insurance hypothesis and to determine the role dispersal timing plays in this response we conducted a field experiment using crustacean zooplankton communities in a subarctic region that is expected to be highly impacted by climate change – Churchill, Canada. Three experiments were conducted where nutrients, salt, and dispersal were manipulated. The three experiments differed in time‐since‐disturbance that the dispersers were added. We found that coarse measures of diversity (i.e. species richness, evenness, and Shannon–Weiner diversity) were generally resistant to large magnitude disturbances, and that dispersal had the most impact on diversity when dispersers were added shortly after disturbance. Ecosystem functioning (chl‐a) was degraded in disturbed communities, but dispersal recovered ecosystem function to undisturbed levels. This spatial insurance for ecosystem function was mediated through changes in community composition and the relative abundance of functional groups. Results suggest that regional diversity and habitat connectivity will be important in the future to maintain ecosystem function by introducing functionally redundant species to promote compensatory dynamics.     

Combined effects of zooplankton grazing and dispersal on the diversity and assembly mechanisms of bacterial metacommunities

Effects of dispersal and the presence of predators on diversity, assembly and functioning of bacterial communities are well studied in isolation. In reality, however, dispersal and trophic interactions act simultaneously and can therefore have combined effects, which are poorly investigated. We performed an experiment with aquatic metacommunities consisting of three environmentally different patches and manipulated dispersal rates among them as well as the presence or absence of the keystone species Daphnia magna. Daphnia magna reduced both local and regional diversity, whereas dispersal increased local diversity but decreased beta‐diversity having no net effect on regional diversity. Dispersal modified the assembly mechanisms of bacterial communities by increasing the degree of determinism. Additionally, the combination of the D. magna and dispersal increased the importance of deterministic processes, presumably because predator‐tolerant taxa were spread in the metacommunity via dispersal. Moreover, the presence of D. magna affected community composition, increased community respiration rates but did not affect bacterial production or abundance, whereas dispersal slightly increased bacterial production. In conclusion, our study suggests that predation by a keystone species such as D. magna and dispersal additively influence bacterial diversity, assembly processes and ecosystem functioning.     

Cooperation‐mediated plasticity in dispersal and colonization

Kin selection theory predicts that costly cooperative behaviors evolve most readily when directed toward kin. Dispersal plays a controversial role in the evolution of cooperation: dispersal decreases local population relatedness and thus opposes the evolution of cooperation, but limited dispersal increases kin competition and can negate the benefits of cooperation. Theoretical work has suggested that plasticity of dispersal, where individuals can adjust their dispersal decisions according to the social context, might help resolve this paradox and promote the evolution of cooperation. Here, we experimentally tested the hypothesis that conditional dispersal decisions are mediated by a cooperative strategy: we quantified the density‐dependent dispersal decisions and subsequent colonization efficiency from single cells or groups of cells among six genetic strains of the unicellular Tetrahymena thermophila that differ in their aggregation level (high, medium, and low), a behavior associated with cooperation strategy. We found that the plastic reaction norms of dispersal rate relative to density differed according to aggregation level: highly aggregative genotypes showed negative density‐dependent dispersal, whereas low‐aggregation genotypes showed maximum dispersal rates at intermediate density, and medium‐aggregation genotypes showed density‐independent dispersal with intermediate dispersal rate. Dispersers from highly aggregative genotypes had specialized long‐distance dispersal phenotypes, contrary to low‐aggregation genotypes; medium‐aggregation genotypes showing intermediate dispersal phenotype. Moreover, highly aggregation genotypes showed evidence for beneficial kin‐cooperation during dispersal. Our experimental results should help to resolve the evolutionary conflict between cooperation and dispersal: cooperative individuals are expected to avoid kin‐competition by dispersing long distances, but maintain the benefits of cooperation by dispersing in small groups.     

Mapping Soil Erosion Prevention Using an Ecosystem Service Modeling Framework for Integrated Land Management and Policy

Current spatially explicit approaches to map and assess ecosystem services are often grounded on unreliable proxy data based on land use/cover to derive ecosystem service indicators. These approaches fail to make a distinction between the actual service provision and the underlying ecosystem capacity to provide the service. We present an integrative conceptual framework to estimate the provision of soil erosion prevention by combining the structural impact of soil erosion and the social–ecological processes that allow for its mitigation. The framework was tested and illustrated in the Portel municipality in Southern Portugal, a Mediterranean silvo-pastoral system that is prone to desertification and soil degradation. The results show a clear difference in the spatial and temporal distribution of the capacity for ecosystem service provision and the actual ecosystem service provision. It also shows that although the average actual ecosystem service provision in the region is sufficient to mitigate the existing structural impact, vulnerable areas can be identified where significant soil losses are not mitigated at present. This becomes more significant when comparing different land management intensities. Considering these results, we argue that the general assumption that there is an almost direct relation between the capacity for ecosystem service provision of a given area and the actual ecosystem service provision is wrong. We also discuss how the framework presented here could be used to support land management and policy, and how it can be adapted for other regulating services.     

Evolution of body condition‐dependent dispersal in metapopulations

Body condition‐dependent dispersal strategies are common in nature. Although it is obvious that environmental constraints may induce a positive relationship between body condition and dispersal, it is not clear whether positive body conditional dispersal strategies may evolve as a strategy in metapopulations. We have developed an individual‐based simulation model to investigate how body condition–dispersal reaction norms evolve in metapopulations that are characterized by different levels of environmental stochasticity and dispersal mortality. In the model, body condition is related to fecundity and determined either by environmental conditions during juvenile development (adult dispersal) or by those experienced by the mother (natal dispersal). Evolutionarily stable reaction norms strongly depend on metapopulation conditions: positive body condition dependency of dispersal evolved in metapopulation conditions with low levels of dispersal mortality and high levels of environmental stochasticity. Negative body condition‐dependent dispersal evolved in metapopulations with high dispersal mortality and low environmental stochasticity. The latter strategy is responsible for higher dispersal rates under kin competition when dispersal decisions are based on body condition reached at the adult life stage. The evolution of both positive and negative body condition‐dependent dispersal strategies is consequently likely in metapopulations and depends on the prevalent environmental conditions.     

The ideal free distribution as an evolutionarily stable strategy

We examine the evolutionary stability of strategies for dispersal in heterogeneous patchy environments or for switching between discrete states (e.g. defended and undefended) in the context of models for population dynamics or species interactions in either continuous or discrete time. There have been a number of theoretical studies that support the view that in spatially heterogeneous but temporally constant environments there will be selection against unconditional, i.e. random, dispersal, but there may be selection for certain types of dispersal that are conditional in the sense that dispersal rates depend on environmental factors. A particular type of dispersal strategy that has been shown to be evolutionarily stable in some settings is balanced dispersal, in which the equilibrium densities of organisms on each patch are the same whether there is dispersal or not. Balanced dispersal leads to a population distribution that is ideal free in the sense that at equilibrium all individuals have the same fitness and there is no net movement of individuals between patches or states. We find that under rather general assumptions about the underlying population dynamics or species interactions, only such ideal free strategies can be evolutionarily stable. Under somewhat more restrictive assumptions (but still in considerable generality), we show that ideal free strategies are indeed evolutionarily stable. Our main mathematical approach is invasibility analysis using methods from the theory of ordinary differential equations and nonnegative matrices. Our analysis unifies and extends previous results on the evolutionary stability of dispersal or state-switching strategies.     

Genomic comparison of chitinolytic enzyme systems from terrestrial and aquatic bacteria

Chitin degradation ability is known for many aquatic and terrestrial bacterial species. However, differences in the composition of chitin resources between aquatic (mainly exoskeletons of crustaceans) and terrestrial (mainly fungal cell walls) habitats may have resulted in adaptation of chitinolytic enzyme systems to the prevalent resources. We screened publicly available terrestrial and aquatic chitinase‐containing bacterial genomes for possible differences in the composition of their chitinolytic enzyme systems. The results show significant differences between terrestrial and aquatic bacterial genomes in the modular composition of chitinases (i.e. presence of different types of carbohydrate binding modules). Terrestrial Actinobacteria appear to be best adapted to use a wide variety of chitin resources as they have the highest number of chitinase genes, the highest diversity of associated carbohydrate‐binding modules and the highest number of CBM33‐type lytic polysaccharide monooxygenases. A ctinobacteria do also have the highest fraction of genomes containing β‐1, 3‐glucanases, enzymes that may reinforce the potential for degrading fungal cell walls. The fraction of bacterial chitinase‐containing genomes encoding polyketide synthases was much higher for terrestrial bacteria than for aquatic ones supporting the idea that the combined production of antibiotics and cell‐wall degrading chitinases can be an important strategy in antagonistic interactions with fungi.     

Negative density-dependent dispersal emerges from the joint evolution of density- and body condition-dependent dispersal strategies

Empirical studies have documented both positive and negative density-dependent dispersal, yet most theoretical models predict positive density dependence as a mechanism to avoid competition. Several hypotheses have been proposed to explain the occurrence of negative density-dependent dispersal, but few of these have been formally modeled. Here, we developed an individual-based model of the evolution of density-dependent dispersal. This model is novel in that it considers the effects of density on dispersal directly, and indirectly through effects on individual condition. Body condition is determined mechanistically, by having juveniles compete for resources in their natal patch. We found that the evolved dispersal strategy was a steep, increasing function of both density and condition. Interestingly, although populations evolved a positive density-dependent dispersal strategy, the simulated metapopulations exhibited negative density-dependent dispersal. This occurred because of the negative relationship between density and body condition: high density sites produced low-condition individuals that lacked the resources required for dispersal. Our model, therefore, generates the novel hypothesis that observed negative density-dependent dispersal can occur when high density limits the ability of organisms to disperse. We suggest that future studies consider how phenotype is linked to the environment when investigating the evolution of dispersal.     

Dispersal network structure and infection mechanism shape diversity in a coevolutionary bacteria-phage system

Resource availability, dispersal and infection genetics all have the potential to fundamentally alter the coevolutionary dynamics of bacteria–bacteriophage interactions. However, it remains unclear how these factors synergise to shape diversity within bacterial populations. We used a combination of laboratory experiments and mathematical modeling to test how the structure of a dispersal network affects host phenotypic diversity in a coevolving bacteria-phage system in communities of differential resource input. Unidirectional dispersal of bacteria and phage from high to low resources consistently increased host diversity compared with a no dispersal regime. Bidirectional dispersal, on the other hand, led to a marked decrease in host diversity. Our mathematical model predicted these opposing outcomes when we incorporated modified gene-for-gene infection genetics. To further test how host diversity depended on the genetic underpinnings of the bacteria-phage interaction, we expanded our mathematical model to include different infection mechanisms. We found that the direction of dispersal had very little impact on bacterial diversity when the bacteria-phage interaction was mediated by matching alleles, gene-for-gene or related infection mechanisms. Our experimental and theoretical results demonstrate that the effects of dispersal on diversity in coevolving host–parasite systems depend on an intricate interplay of the structure of the underlying dispersal network and the specifics of the host–parasite interaction.     

Impact of soil heat on reassembly of bacterial communities in the rhizosphere microbiome and plant disease suppression

The rhizosphere microbiome offers a range of ecosystem services to the plant, including nutrient acquisition and tolerance to (a)biotic stress. Here, analysing the data by Mendes et al. (2011), we show that short heat disturbances (50 or 80 °C, 1 h) of a soil suppressive to the root pathogenic fungus Rhizoctonia solani caused significant increase in alpha diversity of the rhizobacterial community and led to partial or complete loss of disease protection. A reassembly model is proposed where bacterial families that are heat tolerant and have high growth rates significantly increase in relative abundance after heat disturbance, while temperature‐sensitive and slow‐growing bacteria have a disadvantage. The results also pointed to a potential role of slow‐growing, heat‐tolerant bacterial families from Actinobacteria and Acidobacteria phyla in plant disease protection. In conclusion, short heat disturbance of soil results in rearrangement of rhizobacterial communities and this is correlated with changes in the ecosystem service disease suppression.     

Frugivory on Persea lingue in temperate Chilean forests: interactions between fruit availability and habitat fragmentation across multiple spatial scales

Habitat degradation and fragmentation are expected to reduce seed dispersal rates by reducing fruit availability as well as the movement and abundance of frugivores. These deleterious impacts may also interact with each other at different spatial scales, leading to nonlinear effects of fruit abundance on seed dispersal. In this study we assessed whether the degradation and fragmentation of southern Chilean forests had the potential to restrict seed dispersal the lingue (Persea lingue) tree, a fleshy-fruited tree species. Of five frugivore bird species, the austral thrush (Turdus falcklandii) and the fire-eyed diucon (Xolmis pyrope) were the only legitimate seed dispersers as well as being the most abundant species visiting lingue trees. The results showed little or no direct effect of habitat fragmentation on seed dispersal estimates, possibly because the assemblage of frugivore birds was comprised habitat-generalist species. Instead, the number of fruits removed per focal tree exhibited an enhanced response to crop size, but only in the more connected fragments. In the fruit-richer fragment networks, there was an increased fragment-size effect on the proportion of fruits removed in comparison to fruit-poor networks in which the fragment size effect was spurious. We suggest that such nonlinear effects are widespread in fragmented forest regions, resulting from the link between the spatial scales over which frugivores sample resources and the spatial heterogeneity in fruiting resources caused by habitat fragmentation and degradation.     

Urban ecosystem services assessment along a rural–urban gradient: A cross-analysis of European cities

The main objective of this paper is to present an assessment approach for ecosystem services in an urban context covering the local and the regional scale. It was applied to different European cities. A set of indicators representing important urban ecosystem goods and services – local climate regulation, air cooling and recreation – was tested using spatial data along an urban–rural gradient. The results show that there is neither a typical rural–urban gradient in terms of urban ecosystem service provisioning nor a uniform urban spatial pattern of service provisioning that can serve as a generic model for cities. The results demonstrate that (1) core cities do not necessarily provide fewer ecosystem services compared to their regions and (2) there were no patches found within the four case study cities where all of the indicators report very high performance values. The analysis further shows that a high degree of imperviousness does not necessarily entail low ecosystem service provisioning if an urban structure contains a considerable amount of mature trees which support carbon storage and biodiversity. The results of the present paper provide insights into potentials and trade-offs between different urban ecosystem services that should be considered during urban planning when setting targets and establishing thresholds to protect environmental resources, ecosystem services and biodiversity for residents.