The distribution of positive and negative species interactions across environmental gradients on a dual-lattice model

There has been considerable recent interest in understanding the role of positive inter-specific interactions within ecology, and significant progress has been made both empirically and theoretically. Similarly, considerable progress has been made in improving our understanding of the mechanisms that limit species' ranges. In this contribution, we seek to understand the setting of species' borders when some species within the assemblage exhibit positive inter-specific interactions. We use a spatially explicit dual-lattice simulation model to explore the distribution of different interactions across environmental gradients. We first simulate community dynamics when there is either a gradient in reproductive rate or in mortality. We then consider what happens when gradients in reproduction and mortality run in parallel or perpendicular to one another. If the stress gradient impacts on reproductive potential, positive interactions are found where there is high abiotic stress. In this instance, the mutualists are able to tolerate an environment that the cheaters cannot. However, when the stress gradient influences mortality, we find that the mutualists occur as a stripe surrounded by cheaters both towards the better and the harsher ends of the gradient. Previous theory and most empirical evidence tend to indicate that net positive interactions are likely to occur in environments characterized by high abiotic stress. However, evidence from some stress gradients suggests that the distribution of positive and negative interactions can be more complex, with the most stressful environments being occupied by individuals engaging in negative rather than positive interactions. Our results provide a potential theoretical explanation for these recent field observation, and highlight the need for further theoretical and empirical work to better our understanding of how positive and negative interactions act to determine the limits to species' ranges.     

The interplay between species' positive and negative interactions shapes the community biomass–species richness relationship

We used an individual-based spatially-explicit model to assess the role of facilitation and plant strategies in shaping the 'community biomass–species richness' relationship. Facilitation had few impacts on community's richness under both the most benign (high community biomass) and the most severe (low community biomass) environments where its intensity was weak. From medium to high environmental severity, facilitation increased community richness, because all plant strategies were facilitated. In contrast, from low to medium environmental severity facilitation decreased community richness, because only the most competitive species were facilitated, which induced a decrease in the richness of the stress-tolerant species overwhelming the increase in richness of the competitive species. Above all, our simulations show how 'strategy-dependent' interactions among species combine to shape the humped-back biomass–species richness relationship. It also demonstrates that facilitative effects might have long-term negative effects on species richness, which result is not included in current facilitation models.     

Productivity and species richness across an environmental gradient in a fire-dependent ecosystem

The fire-dependent longleaf pine-wiregrass (Pinus palustris Mill.-Aristida beyrichiana Trin. & Rupr.) savannas of the southeastern United States provide a unique opportunity to examine the relationship between productivity and species richness in a natural ecosystem because of the extremely high number of species and their range across a wide ecological amplitude (sandhills to edges of wetlands). We used a natural gradient to examine how plant species richness and plant community structure vary with standing crop biomass (which in this system is proportional to annual net productivity) as a function of soil moisture and nitrogen mineralization rates in a frequently burned longleaf pine-wiregrass savanna. Highest ground cover biomass and highest species richness were found at the same position along the gradient, the wet-mesic sites. Relative differences in species richness among site types were independent of scale, ranging from 0.01 m(2) to 100 m(2). Nitrogen availability was negatively correlated with species richness. Dominance of wiregrass (in terms of biomass) was consistent across the gradient and not correlated with species richness. Regardless of site type, the community structure of the savannas was characterized by many perennial species with infrequent occurrences, a factor in the low temporal heterogeneity (percent similarity between seasons and years) and high within-site spatial heterogeneity (percent dissimilarity of vegetation composition). The coexistence of numerous species is likely due to the high frequency of fire that removes competing hardwood vegetation and litter and to the suite of fire-adapted perennial species that, once established, are able to persist. Our results suggest that soil moisture is an important factor regulating both the number of species present and community production within the defined gradient of this study.     

Shrubs as ecosystem engineers across an environmental gradient: effects on species richness and exotic plant invasion

Ecosystem-engineering plants modify the physical environment and can increase species diversity and exotic species invasion. At the individual level, the effects of ecosystem engineers on other plants often become more positive in stressful environments. In this study, we investigated whether the community-level effects of ecosystem engineers also become stronger in more stressful environments. Using comparative and experimental approaches, we assessed the ability of a native shrub (Ericameria ericoides) to act as an ecosystem engineer across a stress gradient in a coastal dune in northern California, USA. We found increased coarse organic matter and lower wind speeds within shrub patches. Growth of a dominant invasive grass (Bromus diandrus) was facilitated both by aboveground shrub biomass and by growing in soil taken from shrub patches. Experimental removal of shrubs negatively affected species most associated with shrubs and positively affected species most often found outside of shrubs. Counter to the stress-gradient hypothesis, the effects of shrubs on the physical environment and individual plant growth did not increase across the established stress gradient at this site. At the community level, shrub patches increased beta diversity, and contained greater rarified richness and exotic plant cover than shrub-free patches. Shrub effects on rarified richness increased with environmental stress, but effects on exotic cover and beta diversity did not. Our study provides evidence for the community-level effects of shrubs as ecosystem engineers in this system, but shows that these effects do not necessarily become stronger in more stressful environments.     

Structural validation of an individual-based model for plague epidemics simulation

Plague remains endemic in many countries in the world and Madagascar is currently the country where the highest number of human plague cases is reported every year. The investigation of causal factors, which command the disease dynamics in rodent populations, is a crucial step to forecast, control and anticipate the infection extension to humans. This paper presents simulation results obtained from an epidemic model, SIMPEST, designed to simulate bubonic plague in a rodent population at a high level of spatial and temporal resolution. We developed a structurally realistic individual-based model, mobilizing knowledge about fleas and rats behaviour, inter-individual plague transmission, and disease evolution in individual organisms, so that the model reflects the way the real system operates and to generate spatial and temporal patterns of disease spread. To assess the structural validity of our simulations, we perform sensitivity analyses on the initial population size and spatial distribution, and compare our results with theoretical statements, garnered from both previous modelling experiences and repeated field observations. We show our results are consistent with referents about population size conditions for a disease to invade and persist and the effect of the contact network on disease dynamics.     

The interplay between above‐ and below‐ground plant–plant interactions along an environmental gradient: insights from two‐layer zone‐of‐influence models

The changes in plant–plant interactions along environmental gradients have been a focus of recent ecological research. It has been suggested that both above‐ and below‐ground competition and their interplay vary along gradients, but few studies have investigated this idea, and in most cases, the role of facilitation has not been considered, despite its importance in high stress environments. Here we used two‐layer ‘zone‐of‐influence’ models to simulate the effects of facilitation, size‐asymmetry of competition, abiotic stress, resource availability and the balance of root–shoot growth on shoot and root interactions and their interplay along an environmental gradient. In the absence of facilitation, shoot and total competition became weaker, while root competition and the interplay between shoot and root competition were unchanged under increasing stress when root competition was completely symmetric. In contrast, shoot, root, total interactions and the interplay between shoot and root interactions were all negative, and they increased with increasing stress when root competition was size‐symmetric. When facilitation was included in the models, net effects of shoot, root, total interactions and the interplay of root–shoot interactions were very different from those without facilitation, and many were positive under highly stressful conditions. The type of stress (non‐resource or resource) did not significantly influence the simulation results. Our study provides an alternative interpretation of the interplay between above‐ and below‐ground plant–plant interactions across an environmental gradient.     

植物间正相互作用对种群动态和群落结构的影响:基于个体模型的研究进展

植物间的相互作用对种群动态和群落结构有着重要的影响。大量的野外实验已经揭示了正相互作用(互利)在群落中的普遍存在及其重要性。为了弥补野外实验方法的不足, 模型方法被越来越多地应用于正相互作用及其生态学效应的研究中。该文基于个体模型研究, 探讨了植物间正相互作用对种群动态和群落结构的影响。介绍了植物间正相互作用的定义和发生机制、植物间相互作用与环境梯度的关系。正相互作用是指发生在相邻的植物个体之间, 至少对其中一个个体有益的相互作用。植物通过直接(生境改善或资源富集)或间接(协同防御等)作用使局部环境有利于邻体而发生正相互作用。胁迫梯度假说认为互利的强度或重要性随着环境胁迫度的增加而增加, 但是越来越多的经验研究认为胁迫梯度假说需要改进。以网格模型和影响域模型为例, 介绍了基于个体的植物间相互作用模型方法。基于个体模型, 对近年来国内外正相互作用对种群时间动态(如生物量-密度关系)、空间分布格局和群落结构(如群落生物量-物种丰富度关系)影响的研究进行了总结。指出未来的研究应集中在对正相互作用概念和机制的理解, 新的模型, 新的种群、群落, 甚至生态系统问题, 以及在全球变化背景下进行相关的研究。     

Effects of traits,species identity and local environmental conditions on the assessment of interactions: insights from an alpine meadow community

Aims The prediction that facilitation is the dominant interaction in physically stressful conditions has been supported by many but not all field studies. In the present paper, we tested the effects of the identity of species, the local environmental conditions and the currencies of performance measurement on such variation.Methods Using contrasting two plots, six species, and up to five multiple traits, we comprehensively explored the effects of the above factors on the assessment of plant interactions in an alpine meadow of the Qing-Hai Tibetan Plateau. Additionally, we attempted to figure out the possible mechanisms underlying the responses observed. The data were analysed by both standard ANOVAs and multivariate statistics.Important findings Our results demonstrated that the response to the removal of neighbours was both species and trait specific, and the effect of the local environmental conditions was dependent on the species involved. The contrast between plots had crucial influence on the net interactions of Kobresia macrantha, but little effect on Elymus nutans. Regarding the abiotic conditions, neighbours had significant impact on soil temperature, moist and solar radiation. The results contribute to advance our knowledge on the potential underlying factors influencing the assessment of facilitation.     

The complex interplay between mosquito positive and negative regulators of Plasmodium development

The malaria parasite, Plasmodium, requires sexual development in the mosquito before it can be transmitted to the vertebrate host. Mosquito genes are able to substantially modulate this process, which can result in major decreases in parasite numbers. Even in susceptible mosquitoes, haemolymph proteins implicated in systemic immune reactions, together with local epithelial responses, cause lysis of more than 80% of the ookinetes that cross the mosquito midgut. In a refractory mosquito strain, immune responses lead to melanisation of virtually all parasites. Conversely, certain mosquito genes have an opposite effect: they are used by the parasite to evade defence reactions. Detailed understanding of the interplay between positive and negative regulators of parasite development could lead to the generation of novel approaches for malaria control through the vector.     

Membrane remodeling from N-BAR domain interactions: insights from multi-scale simulation

Liposome remodeling processes (e.g., vesiculation and tubulation) due to N-BAR domain interactions with the lipid bilayer are explored with a multi-scale simulation approach. Results from atomistic-level molecular dynamics simulations of membrane binding to the concave face of N-BAR domains are used along with discretized mesoscopic field-theoretic simulations to examine how the spontaneous curvature fields generated by N-BAR domains result in membrane remodeling. It is found that tubulation can be generated by anisotropic N-BAR spontaneous curvature fields, whereas vesiculation is only observed with isotropic N-BAR spontaneous curvature fields at high density. The results of the multi-scale simulations provide insight into recent experimental observations.     

General response patterns of fish populations to stress: an evaluation using an individual-based simulation model

General response patterns of fish populations tostress, originally proposed by Colby for fisheriesrehabilitation and later adapted by Munkittrick forcontaminants, were evaluated using an individual-basedsimulation model. General response patterns relatechanges in population-level variables to the type ofstress. The model follows the daily growth,mortality, and spawning of individual yellow perch andwalleye through their lifetime, and was corroboratedusing Oneida Lake data. Two versions of the model wereused: population (yellow perch only) and community(dynamic predation on yellow perch by walleye). Eightstresses were imposed on the population and communityversions of the model and 100-year simulations wereperformed. Response patterns were defined by changesin predicted yellow perch mean population abundance,mean age of adults, and mean adult growth (representedby mean length at age-7). Proposed response patternswere similar to those predicted using the populationversion of the model. Simulations using the communityversion of the model distorted the response patterns,either causing amplification, dampening, or reversalof many of the patterns. Predicted response patternsbecame unique when additional variables were included.Our model results suggest that caution is appropriatein interpreting general response patterns based onmean age, or when the population of interest plays amajor role in a relatively simple food web. The responsepattern approach may be better at identifying the lifestage impacted rather than the mechanism of the stress.     

Modelling interactions between farmer practices and fattening pig performances with an individual-based model

European pig production continues to encounter economic and environmental challenges. To address these issues, methods have been developed to assess performances of pig production systems. Recent studies indicate that considering variability in performances among pigs improves the accuracy and reliability of results compared with modelling an average animal. Our objective was to develop a pig fattening unit model able to (i) simulate individual pig performances, including their variability in interaction with farmers’ practices and management, and (ii) assess their effects on technical, economic and environmental performances. Farmer practices included in the model were chosen from a typology generated from on-farm surveys focused on batch management, pig allocation to pens, pig feeding practices, practices of shipping to the slaughterhouse, and management of the remaining pigs. Pigs are represented using an individual-based model adapted from the InraPorc® model. To illustrate the model’s abilities, four scenarios were simulated that combine two feed rationing plans (ad libitum, restricted to 2.5 kg/day) and two feed sequence plans (two-phase, 10-phase). Analysis of variance was performed on the simulated technical, economic and environmental indicators (calculated via Life Cycle Assessment). The feed rationing plan and feed sequence plan significantly affected all indicators except for the premium per pig, for which the feed sequence plan did not have a significant effect. The ‘restricted 10-phase’ scenario maximised gross margin of the fattening unit (14.2 €/pig) and minimised environmental impacts per kg of pig produced. In contrast, the ‘ad libitum two-phase’ scenario generated the lowest margin (8.20 €/pig) and the highest environmental impacts. The model appears to be a promising tool to assess effects of farmers’ practices, pig characteristics and farm infrastructure on technical, economic and environmental performances of the fattening unit, and to investigate the potential of improvement. However, further work is needed, based on virtual experiments, in order to evaluate the effects of a larger diversity of practices.     

Local coexistence of tree species and the dynamics of global distribution pattern along an environmental gradient: a simulation study

A spatially explicit tree-based model was used to demonstrate the effects of a mechanism promoting multiple-species coexistence on the development of vegetation zonation and its response to climate change. Temporal fluctuation in reproduction was incorporated as the mechanism, which facilitates the persistence of less competitive species. Four hypothetical tree species with different temperature dependencies of seed production were randomly located over a landscape represented by 2,000×40 cells. Each cell can sustain a single tree at most. A zonal distribution pattern emerged corresponding to the temperature gradient along the long axis of the landscape. When there was a temporal variation in seed production, species became distributed over a wider range than that when seed production was constant. When the whole landscape was warmed, the distribution range of each species shifted towards the cool end of the landscape. However, the migration was retarded due to competition for vacant spaces with the remnant species which had dominated the location before the warming. Temporal fluctuation in reproduction facilitated the migration because it enhanced the persistence of minority species and, thus, the invasion and establishment of new species in the area dominated by other species.     

A new model of myofibroblast-cardiomyocyte interactions and their differences across species

Although coupling between cardiomyocytes and myofibroblasts is well known to affect the physiology and pathophysiology of cardiac tissues across species, relating these observations to humans is challenging because the effect of this coupling varies across species and because the sources of these effects are not known. To identify the sources of cross-species variation, we built upon previous mathematical models of myofibroblast electrophysiology and developed a mechanoelectrical model of cardiomyocyte-myofibroblast interactions as mediated by electrotonic coupling and transforming growth factor-β1. The model, as verified by experimental data from the literature, predicted that both electrotonic coupling and transforming growth factor-β1 interaction between myocytes and myofibroblast prolonged action potential in rat myocytes but shortened action potential in human myocytes. This variance could be explained by differences in the transient outward K⁺ current associated with differential Kv4.2 gene expression across species. Results are useful for efforts to extrapolate the results of animal models to the predicted effects in humans and point to potential therapeutic targets for fibrotic cardiomyopathy.     

Stochastic simulation of size variation in turbot: possible causes analysed with an individual-based model

Causes of size variation in a population of juvenile turbot were studied using an individual based model (IBM). Each simulation started with 800 (divided into eight groups of 100 each) 120-day-old (posthatch) juveniles and was run for 140 days, and the data gained from model simulations compared directly with the result of a laboratory study with size-graded turbot. Stochastic growth with memory, which was included in the models as an individual genetical growth rate variation, is important in explaining size variation, and the combination between individual genetic growth rate and social interactions related to size-dependent hierarchies also contributes to size variation. The use of size-dependent growth rate alone fails to explain size variation, and is of little value in predicting size variation in turbot culture. Further, the results indicate formation of different types of size hierarchies for different sizes of juvenile turbot.     

Individual variability and population regulation: an individual-based model

To study the influence of individual variability on population dynamics an individual-based model of the dynamics of a single population consisting of different individuals is constructed. The model is based on differences in individual assimilation rates due to intraspecific competition and variability of initial weights. The model exhibits "imperfect regulation", i.e., the number of individuals in the population oscillates and sooner or later the population becomes extinct. When individual variability is included, the model produces longer population extinction times than without individual variability. The average extinction time is not however a monotonic function of the degree of individual variability.     

Ontogenetic and individual diet variation in amphibian larvae across an environmental gradient

1. Variation among individuals within size or age classes can have profound effects on community dynamics and food‐web structure. We investigated the potential influence of habitat disturbance on intrapopulation niche variation. 2. Amphibians occupy a range of lentic habitats from short‐hydroperiod intermittent ponds to long‐hydroperiod permanent ponds. We quantified ontogenetic diet variation and individual specialisation in wood frog tadpoles (Lithobates sylvaticus) and blue‐spotted salamander larvae (Ambystoma laterale) to investigate the influence of hydroperiod on population niche width across a natural hydroperiod gradient using stable isotope and gut content analyses. In one of the few tests using larval forms, we tested the niche variation hypothesis, which predicts that populations with larger niche widths also have increased individual variation. 3. Our results support the niche variation hypothesis, indicating that more generalised populations exhibit higher within‐individual diet variation. We report gradual changes in the relative importance of diet items, decreased dietary overlap and increased trophic position in L. sylvaticus throughout development. A. laterale became more enriched in δ¹³C and increased in δ¹⁵N throughout its larval period. We did not find a relationship between hydroperiod and niche parameters, indicating that niches are conserved across heterogeneous habitats. In contrast to most documented cases, we estimated low levels of individual specialisation in amphibian larvae. 4. Amphibians are an important link between aquatic and terrestrial ecosystems, whereby diet shifts can influence food‐web structure by altering energy flow pathways and the trophic position of higher consumers, ultimately changing food‐chain length.     

Trophic omnivory across a productivity gradient: intraguild predation theory and the structure and strength of species interactions

Intraguild predation theory centres on two predictions: (i) for an omnivore and an intermediate predator (IG-prey) to coexist on shared resources, the IG-prey must be the superior resource competitor, and (ii) increasing resource productivity causes the IG-prey''s equilibrium abundance to decline. I tested these predictions with a series of species-rich food webs along New Zealand''s rocky shores, focusing on two predatory whelks, Haustrum haustorium, a trophic omnivore, and Haustrum scobina, the IG-prey. In contrast to theory, the IG-prey''s abundance increased with productivity. Furthermore, feeding rates and allometric considerations indicate a competitive advantage for the omnivore when non-shared prey are considered, despite the IG-prey''s superiority for shared prey. Nevertheless, clear and regular cross-gradient changes in network structure and interaction strengths were observed that challenge the assumptions of current theory. These insights suggest that the consideration of consumer-dependent functional responses, non-equilibrium dynamics, the dynamic nature of prey choice and non-trophic interactions among basal prey will be fruitful avenues for theoretical development.     

The evolution of species interactions across natural landscapes

Given the potential for rapid and microgeographical adaptation, ecologists increasingly are exploring evolutionary explanations for community patterns. Biotic selection can generate local adaptations that alter species interactions. Although some gene flow might be necessary to fuel local adaptation, higher gene flow can homogenise traits across regions and generate local maladaptation. Herein, I estimate the contributions of local biotic selection, gene flow and spatially autocorrelated biotic selection to among-population divergence in traits involved in species interactions across 75 studies. Local biotic selection explained 6.9% of inter-population trait divergence, an indirect estimate of restricted gene flow explained 0.1%, and spatially autocorrelated selection explained 9.3%. Together, biotic selection explained 16% of the variance in population trait means. Most biotic selection regimes were spatially autocorrelated. Hence, most populations receive gene flow from populations facing similar selection, which could allow for local adaptation despite moderate gene flow. Gene flow constrained adaptation in studies conducted at finer spatial scales as expected, but this effect was often confounded with spatially autocorrelated selection. Results indicate that traits involved in species interactions might often evolve across landscapes, especially when biotic selection is spatially autocorrelated. The frequent evolution of species interactions suggests that evolutionary processes might often influence community ecology.