Developing an ecological context for allelopathy

There has been a renewed interest in allelopathy as a plant–plant interaction as more plant ecologists have become involved in studying biological invasions. This resurgence highlights a major deficiency in our understanding of allelopathy—the lack of a well-developed ecological context for the interaction. In contrast to allelopathy, the plant–plant interaction of competition has a strong theoretical foundation as well as a large body of supporting empirical studies. We suggest that the plant-herbivore defense literature provides a mature and well-developed framework from which a broader ecological context for allelopathy can be developed. Here, we discuss three broad classes of questions, drawn from the herbivore defense literature, which may help to develop an appropriate ecological context for allelopathy. These questions focus on (1) variation in allelopathic expression within species, (2) community level variation in allelopathy across species, and (3) variation in the impacts of allelopathy on associated species. Addressing such broad population and community level themes in a variety of systems will be necessary to fully develop an ecological context for allelopathy and provide a theoretical basis for understanding its role in plant communities.     

Allelopathic adaptation can cause competitive coexistence

The maintenance of plant diversity is often explained by the ecological and evolutionary consequences of resource competition. Recently, the importance of allelopathy for competitive interactions has been recognized. In spite of such interest in allelopathy, we have few theories for understanding how the allelopathy influences the ecological and evolutionary dynamics of competing species. Here, I study the coevolutionary dynamics of two competing species with allelopathy in an interspecific competition system, and show that adaptive trait dynamics can cause cyclic coexistence. In addition, very fast adaptation such as phenotypic plasticity is likely to stabilize the population cycles. The results suggest that adaptive changes in allelopathy can lead to cyclic coexistence of plant species even when their ecological characters are very similar and interspecific competition is stronger than intraspecific competition, which should destroy competitive coexistence in the absence of adaptation.     

Functional correlates of allelopathic potential in a successional plant community

Allelopathy, plant–plant interactions mediated through chemical production, is an active area of ecological research. Despite this widespread interest, we still lack community scale information on the prevalence of this interaction and the types of species that may be expected to be allelopathic. To address this research need, the allelopathic potential of 65 plant species from all stages of succession in the Piedmont region of New Jersey, USA, was determined with laboratory bioassays. The strength of each species’ allelopathic activity was then related to life form, origin, and fundamental plant traits. The vast majority of species tested exhibited significant allelopathic effects in the bioassays, with many of these having fairly strong effects. Overall, the allelopathic potential of species decreased with life span, roughly following the successional transitions from short-lived to long-lived herbs and to woody species. Herbaceous species on average were more allelopathic than woody species, but there was no difference between native and non-native species once life form was accounted for. In a principal components analysis, allelopathy was associated with other plant traits, but these relationships differed between woody and herbaceous species. Allelopathic potential was positively associated with plant height in herbaceous species, but negatively associated with height, leaf mass, and seed mass in woody species. These results indicate that allelopathy may be a quite common ecological strategy in plants and is equally common in both native and non-native species. The linkage of allelopathy with other plant functional traits suggests that allelopathy can and should be integrated into the broader suite of plant strategies that are studied.     

Drivers of plant individual-based pollinator visitation network topology in an arid ecosystem

Interactions with pollinators underlie the structure and function of plant communities. Network analysis is a valuable tool for studying plant-pollinator interactions, but these networks are most frequently built by aggregating interactions at the species level. Interactions are between individuals and an advantage of individual-based networks is the ability to integrate inter-individual variation in traits and environmental context within complex ecological networks. We studied the influence of inter-individual variation on pollinator sharing among foundation shrubs and cactus in a desert ecosystem using plant individual-based pollinator visitation networks. We hypothesized that the traits that alter attractiveness of plants to pollinators will also influence an individual plant's role within the visitation network. Foundation plants growing with higher densities of nearby blooming shrubs had higher pollinator visitation rates and had greater access to the conspecific mating pool, suggesting widespread and diffuse pollination facilitation within this community. Further, shrub density influenced the role of betweenness centrality and the effective number of partners (e^H). Floral display size also influenced the effective number of interaction partners but did not directly influence the centrality measures for individual plants or other measures of network structure despite increasing visitation rates. The individual-based visitation networks were significantly modular and module membership was predicted by species identity and pollinator visitation rates. Ecological and individual context mediate the outcome of pollinator-mediated interactions and are fundamental drivers of whole community structure. This study shows that the density of immediate neighbours can influence the overall structure of plant-pollinator interaction networks. Exploring the contribution of intraspecific variation to community interaction networks will improve our understanding of drivers of community-level ecological dynamics.     

Challenges,achievements and opportunities in allelopathy research

Abstract

Allelopathy is defined as the suppression of any aspect of growth and/or development of one plant by another through the release of chemical compounds. Although allelopathic interference has been demonstrated many times using in vitro experiments, few studies have clearly demonstrated allelopathy in natural settings. This difficulty reflects the complexity in examining and demonstrating allelopathic interactions under field conditions. In this paper we address a number of issues related to the complexity of allelopathic interference in higher plants: These are: (i) is a demonstrated pattern or zone of inhibition important in documenting allelopathy? (ii) is it ecologically relevant to explain the allelopathic potential of a species based on a single bioactive chemical? (iii) what is the significance of the various modes of allelochemical release from the plant into the environment? (iv) do soil characteristics clearly influence allelopathic activity? (v) is it necessary to exclude other plant interference mechanisms?, and (vi) how can new achievements in allelopathy research aid in solving problems related to relevant ecological issues encountered in research conducted upon natural systems and agroecosystems? A greater knowledge of plant interactions in ecologically relevant environments, as well as the study of biochemical pathways, will enhance our understanding of the role of allelopathy in agricultural and natural settings. In addition, novel findings related to the relevant enzymes and genes involved in production of putative allelochemicals, allelochemical persistence in the rhizosphere, the molecular target sites of allelochemicals in sensitive plant species and the influence of allelochemicals upon other organisms will likely lead to enhanced utilization of natural products for pest management or as pharmaceuticals and nutraceuticals. This review will address these recent findings, as well as the major challenges which continue to influence the outcomes of allelopathy research.     

The future of plant-pollinator diversity: understanding interaction networks across time, space, and global change

Structural analysis of plant-pollinator networks has revealed remarkably high species and interaction diversity and highlighted the species important for pollination services. Although techniques to analyze plant-pollinator networks began to emerge a decade ago, the characterization of spatiotemporal variation of interactions is still in its infancy. Understanding the ecological and evolutionary causes and consequences of spatial and temporal variation in plant-pollinator interactions is important for both basic and applied questions in community structure and function, the evolution of floral traits, and the development of optimal conservation strategies. Here we review observational, theoretical, and experimental studies of temporal and spatial variation in plant-pollinator interaction networks to establish a foundation for future studies to incorporate perspectives in spatiotemporal variation. Such perspectives are crucial given the rapid environmental changes associated with habitat loss, climate change, and biological invasions, which we discuss in this context. The inherent plasticity of plant-pollinator interactions and network structure suggests that many species should be able to persist by responding to environmental changes quickly, even though the identity of their mutualistic partners may change.     

Integrating theory into disturbance interaction experiments to better inform ecosystem management

Managing multiple, interacting disturbances is a key challenge to biodiversity conservation, and one that will only increase as global change drivers continue to alter disturbance regimes. Theoretical studies have highlighted the importance of a mechanistic understanding of stressor interactions for improving the prediction and management of interactive effects. However, many conservation studies are not designed or interpreted in the context of theory and instead focus on case‐specific management questions. This is a problem as it means that few studies test the relationships highlighted in theoretical models as being important for ecological management. We explore the extent of this problem among studies of interacting disturbances by reviewing recent experimental studies of the interaction between fire and grazing in terrestrial ecosystems. Interactions between fire and grazing can occur via a number of pathways; one disturbance can modify the other's likelihood, intensity or spatial distribution, or one disturbance can alter the other's impacts on individual organisms. The strength of such interactions will vary depending on disturbance attributes (e.g. size or intensity), and this variation is likely to be nonlinear. We show that few experiments testing fire–grazing interactions are able to identify the mechanistic pathway driving an observed interaction, and most are unable to detect nonlinear effects. We demonstrate how these limitations compromise the ability of experimental studies to effectively inform ecological management. We propose a series of adjustments to the design of disturbance interaction experiments that would enable tests of key theoretical pathways and provide the deeper ecological understanding necessary for effective management. Such considerations are relevant to studies of a broad range of ecological interactions and are critical to informing the management of disturbance regimes in the context of accelerating global change.     

Plant chemical defense allocation constrains evolution of tolerance to community change across a range boundary

Because transplant experiments show that performance usually decreases across species range boundaries, some range limits might develop from factors and processes that prevent adaptation to stressful environments. Here, we determined whether an ecological cost of plant defense involving stress associated with changes in the local plant community may contribute to range limit development in the upland mustard species Boechera stricta. In a common garden experiment of 499 B. stricta plants, performance decreased and a multivariate axis of community structure increased across the boundary, indicating increased stress associated with the community change. There was also significant genetic variation (evolutionary potential) among marker‐inferred inbred lines of B. stricta for tolerance to the stress; however, lines with high basal levels of glucosinolate toxins had lower tolerance to the change in community structure. We suggest that defense allocation, which is also needed across the range, may impede adaptation to the stress associated with the community change and thus contribute to range limit development.     

Allelopathy and exotic plant invasion

The primary hypothesis for the astonishing success of many exotics as community invaders relative to their importance in their native communities is that they have escaped the natural enemies that control their population growth – the `natural enemies hypothesis'. However, the frequent failure of introduced biocontrols, weak consumer effects on the growth and reproduction of some invaders, and the lack of consistent strong top-down regulation in many natural ecological systems indicate that other mechanisms must be involved in the success of some exotic plants. One mechanism may be the release by the invader of chemical compounds that have harmful effects on the members of the recipient plant community (i.e., allelopathy). Here, we provide an abbreviated compilation of evidence for allelopathy in general, present a detailed case study for Centaurea diffusa, an invasive Eurasian forb in western North America, and review general evidence for allelopathic effects of invasive plants in native communities. The primary rationale for considering allelopathy as a mechanism for the success of invaders is based on two premises. First, invaders often establish virtual monocultures where diverse communities once flourished, a phenomenon unusual in natural communities. Second, allelopathy may be more important in recipient than in origin communities because the former are more likely to be naïve to the chemicals possessed by newly arrived species. Indeed, results from experiments on C. diffusa suggest that this invader produces chemicals that long-term and familiar Eurasian neighbors have adapted to, but that C. diffusa's new North American neighbors have not. A large number of early studies demonstrated strong potential allelopathic effects of exotic invasive plants; however, most of this work rests on controversial methodology. Nevertheless, during the last 15 years, methodological approaches have improved. Allelopathic effects have been tested on native species, allelochemicals have been tested in varying resource conditions, models have been used to estimate comparisons of resource and allelopathic effects, and experimental techniques have been used to ameliorate chemical effects. We do not recommend allelopathy as a `unifying theory' for plant interactions, nor do we espouse the view that allelopathy is the dominant way that plants interact, but we argue that non-resource mechanisms should be returned to the discussion table as a potential mechanism for explaining the remarkable success of some invasive species. Ecologists should consider the possibility that resource and non-resource mechanisms may work simultaneously, but vary in their relative importance depending on the ecological context in which they are studied. One such context might be exotic plant invasion.     

Intraspecific variability improves environmental matching,but does not increase ecological breadth along a wet‐to‐dry ecotone

It is widely assumed that higher levels of intraspecific variability in one or more traits should allow species to persist under a wider range of environmental conditions. However, few studies have examined whether species that exhibit high variability are found in a wider range of environmental conditions, and whether variability increases the ability of a species to adapt to prevailing ecological gradients. We used four plant functional traits, specific leaf area (SLA), leaf dry matter content (LDMC), leaf carbon to nitrogen ratio (C:N) and maximum plant height in 49 species across a strong environmental gradient to answer three questions: 1) is there evidence for ‘high‐variability’ species (that is, species which show high variability in multiple traits, simultaneously)? 2) are species with more variable traits present across a wider range of environmental conditions than less variable species? And 3) whether more variable species show better trait–environment matching to the prevailing abiotic (soil moisture) gradient at the site? We found little evidence for a ‘high‐variability’ species. Variability was correlated for two leaf traits, SLA and LDMC, while variability in leaf traits and plant height were not correlated. We found little evidence that more variable species were present in more diverse conditions: only variation in SLA was correlated with a wider ecological niche breadth. For plant traits along the soil‐moisture gradient, higher variability led to better trait–environment matching in half of measured traits. Overall, we found little support for the existence of ‘high‐variability’ species, but that variability in SLA is correlated with a wider ecological breadth. We also found evidence that variation in traits can improve trait–environment matching, a relationship which may facilitate our understanding ecological breadth along prevailing gradients, and community assembly on the basis of traits.     

Flavonoids:New Roles for Old Molecules

Flavonoids are ubiquitous in the plant kingdom and have many diverse functions including defense, UV protection, auxin transport inhibition, allelopathy, and flower coloring. Interestingly, these compounds also have considerable biological activity in plant, animal and bacterial systems – such broad activity is accomplished by few compounds. Yet, for all the research over the last three decades, many of the cellular targets of these secondary metabolites are unknown. The many mutants available in model plant species such as Arabidopsis thaliana and Medicago truncatula are enabling the intricacies of the physiology of these compounds to be deduced. In the present review, we cover recent advances in flavonoid research, discuss deficiencies in our understanding of the physiological processes, and suggest approaches to identify the cellular targets of flavonoids.     

Bringing Elton and Grinnell together: a quantitative framework to represent the biogeography of ecological interaction networks

Biogeography has traditionally focused on the spatial distribution and abundance of species. Both are driven by the way species interact with one another, but only recently community ecologists realized the need to document their spatial and temporal variation. Here, we call for an integrated approach, adopting the view that community structure is best represented as a network of ecological interactions, and show how it translates to biogeography questions. We propose that the ecological niche should encompass the effect of the environment on species distribution (the Grinnellian dimension of the niche) and on the ecological interactions among them (the Eltonian dimension). Starting from this concept, we develop a quantitative theory to explain turnover of interactions in space and time – i.e. a novel approach to interaction distribution modeling. We apply this framework to host–parasite interactions across Europe and find that two aspects of the environment (temperature and precipitation) exert a strong imprint on species co‐occurrence, but not on species interactions. Even where species co‐occur, interaction proves to be stochastic rather than deterministic, adding to variation in realized network structure. We also find that a large majority of host‐parasite pairs are never found together, thus precluding any inferences regarding their probability to interact. This first attempt to explain variation of network structure at large spatial scales opens new perspectives at the interface of species distribution modeling and community ecology.     

The relative of species pools in determining plant species richness: an alternative explanation of species coexistence?

Explanations of the pattern of species have traditionally relied on small-scale, local processes occurring in ecological time. Differences in species richness have associated with different mechanisms avoiding competition, such as spatiotemporal heterogeneity (weaker competitors may find a more favourable place or time) or environmental stress (competition is assumed to be less intensive under difficult conditions). More recently, large-scale process have been taken into account, raising such questions as: which plant species may potentially grow in a certain community? Are evolutionary processes and species dispersal responsible for the differences between communities? The species-pool theory attempts to answer these general questions, and information about species pools is needed for the design of experiments where the number of species in a community is manipulated.     

Richness, diversity, and rate of primary succession over 20?year in tropical coastal dunes

The tropical coastal dunes in central Gulf of Mexico have been stabilizing over the last decades resulting in reduced substrate mobility, and promoting primary succession. We describe changes in species richness and diversity in dune vegetation during 20?years. Our questions: (a) Do species richness and diversity increase over time as predicted by models of ecological succession or do they show a hump-backed manner similar to the observations in temperate coastal dunes?, (b) What is the interaction between vegetation cover and diversity and species richness?, (c) Is there a relationship between species diversity and succession rate and does succession rate change over time?, and (d) How do plant functional types change during succession? In order to answer these questions, we set 140 4?×?4?m permanent plots in a mobile dune area and monitored vegetation cover and species richness from 1991 to 2011. In time, diversity increased in a logistic manner toward an asymptotic value once vegetation cover surpassed 60?%. Species richness increased in a humped-back shape, also reaching a maximum peak at 60?% vegetation cover. The succession rate of diversity was measured by the Euclidean distance, and showed a significant humped-back relation, meaning that it was slower in early and late successional stages. The study supports the intermediate disturbance theory. The conservation of coastal dunes vegetation should focus on all, species-poor and species-rich habitats that help to maintain the ecological integrity of these ecosystems. The understanding of community dynamics and diversity patterns becomes an essential component of coastal dune management and conservation.     

Remotely sensed vegetation phenology and productivity along a climatic gradient: on the value of incorporating the dimension of woody plant cover

Aim Woody plants affect vegetation–environment interactions by modifying microclimate, soil moisture dynamics and carbon cycling. In examining broad‐scale patterns in terrestrial vegetation dynamics, explicit consideration of variation in the amount of woody plant cover could provide additional explanatory power that might not be available when only considering landscape‐scale climate patterns or specific vegetation assemblages. Here we evaluate the interactive influence of woody plant cover on remotely sensed vegetation dynamics across a climatic gradient along a sky island. Location The Santa Rita Mountains, Arizona, USA. Methods Using a satellite‐measured normalized difference vegetation index (NDVI) from 2000 to 2008, we conducted time‐series and regression analyses to explain the variation in functional attributes of vegetation (productivity, seasonality and phenology) related to: (1) vegetation community, (2) elevation as a proxy for climate, and (3) woody plant cover, given the effects of the other environmental variables, as an additional ecological dimension that reflects potential vegetation–environment feedbacks at the local scale. Results NDVI metrics were well explained by interactions among elevation, vegetation community and woody plant cover. After accounting for elevation and vegetation community, woody plant cover explained up to 67% of variation in NDVI metrics and, notably, clarified elevation‐ and community‐specific patterns of vegetation dynamics across the gradient. Main conclusions In addition to the environmental factors usually considered – climate, reflecting resources and constraints, and vegetation community, reflecting species composition and relative dominance – woody plant cover, a broad‐scale proxy of many vegetation–environment interactions, represents an ecological dimension that provides additional process‐related understanding of landscape‐scale patterns of vegetation function.     

Linking community and disease ecology: the impact of biodiversity on pathogen transmission

The increasing number of zoonotic diseases spilling over from a range of wild animal species represents a particular concern for public health, especially in light of the current dramatic trend of biodiversity loss. To understand the ecology of these multi-host pathogens and their response to environmental degradation and species extinctions, it is necessary to develop a theoretical framework that takes into account realistic community assemblages. Here, we present a multi-host species epidemiological model that includes empirically determined patterns of diversity and composition derived from community ecology studies. We use this framework to study the interaction between wildlife diversity and directly transmitted pathogen dynamics. First, we demonstrate that variability in community composition does not affect significantly the intensity of pathogen transmission. We also show that the consequences of community diversity can differentially impact the prevalence of pathogens and the number of infectious individuals. Finally, we show that ecological interactions among host species have a weaker influence on pathogen circulation than inter-species transmission rates. We conclude that integration of a community perspective to study wildlife pathogens is crucial, especially in the context of understanding and predicting infectious disease emergence events.     

植物化感物质及化感潜力与土壤养分的相互影响

植物化感作用与许多生态因子有关.土壤养分缺乏,影响着许多植物化感物质的产生,从而影响植物的化感潜力;反过来,植物化感物质也通过络合、吸附、酸溶解、竞争、抑制等方式影响土壤的养分形态和水平.本文总结了植物化感物质及化感潜力与土壤养分的相互影响,并提出了今后该领域值得进一步研究的问题.包括以下几方面：加强植物化感研究与土壤 植物营养学研究的结合,以更深入地阐明植物化感物质、化感作用与土壤养分变化的关系;加强植物化感研究与生态系统养分循环研究的结合,以类似自然（nature-like）的方式模拟自然界植物所受的养分干,使养分干扰的化感研究结果更加逼真、可靠;加强对养分过量及受污染时植物化感作用的研究,为揭示农业和林业生产中植物的相互作用机制和生物量变化机制提供新的思路,为生态保护提供科学依据.     

Soil microbial communities alter allelopathic competition between <Emphasis Type="Italic">Alliaria petiolata</Emphasis> and a native species

Allelopathy has been increasingly invoked as a mechanism facilitating exotic plant invasions. However, studies even on the same target species often yield varying results concerning the strength and importance of allelopathic inhibition, suggesting that the process may depend on the specific environmental context. Here I studied how the allelopathic inhibition of sycamore (Platanus occidentalis) seedlings by garlic mustard (Alliaria petiolata) depended on the presence of a soil microbial community. Using three analytical approaches to quantifying allelopathy, I consistently found allelopathic inhibition only in sterilized soils, suggesting that certain microbial taxa inhibit the process, possibly by degrading the allelochemicals. Determining the environmental contexts that reduce or eliminate allelopathic inhibition could lead to a greater understanding of the spatial variation in invasion success and potentially lead to new avenues for management.     

How do pollinator visitation rate and seed set relate to species’ floral traits and community context?

Differences among plant species in visitation rate and seed set within a community may be explained both by the species’ floral traits and the community context. Additionally, the importance of species’ floral traits vs. community context on visitation rate and seed set may vary among communities. In communities where the pollinator-to-flower ratio is low, floral traits may be more important than community context, as pollinators may have the opportunity to be choosier when visiting plant species. In this study we investigated whether species’ floral traits (flower shape, size and number, and flowering duration) and community context (conspecific and heterospecific flower density, and pollinator abundance) could explain among-species variation in visitation rate and seed set. For this, we used data on 47 plant species from two Norwegian plant communities differing in pollinator-to-flower ratio. Differences among species in visitation rate and seed set within a community could be explained by similar variables as those explaining visitation rate and seed set within species. As expected, we found floral traits to be more important than community context in the community with a lower pollinator-to-flower ratio; whereas in the community with a higher pollinator-to-flower ratio, community context played a bigger role. Our study gives significant insights into the relative importance of floral traits on species’ visitation rate and seed set, and contributes to our understanding of the role of the community context on the fitness of plant species.     

Ant-Plant Interaction in a Tropical Savanna: May the Network Structure Vary over Time and Influence on the Outcomes of Associations?

Plant-animal interactions occur in a community context of dynamic and complex ecological interactive networks. The understanding of who interacts with whom is a basic information, but the outcomes of interactions among associates are fundamental to draw valid conclusions about the functional structure of the network. Ecological networks studies in general gave little importance to know the true outcomes of interactions and how they may change over time. We evaluate the dynamic of an interaction network between ants and plants with extrafloral nectaries, by verifying the temporal variation in structure and outcomes of mutualism for the plant community (leaf herbivory). To reach this goal, we used two tools: bipartite network analysis and experimental manipulation. The networks exhibited the same general pattern as other mutualistic networks: nestedness, asymmetry and low specialization and this pattern was maintained over time, but with internal changes (species degree, connectance and ant abundance). These changes influenced the protection effectiveness of plants by ants, which varied over time. Our study shows that interaction networks between ants and plants are dynamic over time, and that these alterations affect the outcomes of mutualisms. In addition, our study proposes that the set of single systems that shape ecological networks can be manipulated for a greater understanding of the entire system.