Search

How is trait diversity in a community apportioned between and within coevolving species? Disruptive selection may result in either a few species with large intraspecific trait variation (ITV) or many species with different mean traits but little ITV. Similar questions arise in spatially structured communities: heterogeneous environments could result in either a few species that exhibit local adaptation or many species with different mean traits but little local adaptation. To date, theory has been well-equipped to either include ITV or to dynamically determine the number of coexisting species, but not both. Here, we devise a theoretical framework that combines these facets and apply it to the above questions of how trait variation is apportioned within and between species in unstructured and structured populations, using two simple models of Lotka-Volterra competition. For unstructured communities, we find that as the breadth of the resource spectrum increases, ITV goes from being unimportant to crucial for characterizing the community. For spatially structured communities on two patches, we find no local adaptation, symmetric local adaptation, or asymmetric local adaptation, depending on how much the patches differ. Our framework provides a general approach to incorporate ITV in models of eco-evolutionary community assembly. [ABSTRACT FROM AUTHOR]

The spread of an enteric pathogen in the human gut depends on many interacting factors, including pathogen exposure, diet, host gut environment, and host microbiota, but how these factors jointly influence infection outcomes remains poorly characterized. Here we develop a model of host-mediated resource competition between mutualistic and pathogenic taxa in the gut that aims to explain why similar hosts, exposed to the same pathogen, can have such different infection outcomes. Our model successfully reproduces several empirically observed phenomena related to transitions between healthy and infected states, including (1) the nonlinear relationship between pathogen inoculum size and infection persistence, (2) the elevated risk of chronic infection during or after treatment with broad-spectrum antibiotics, (3) the resolution of gut dysbiosis with fecal microbiota transplants, and (4) the potential protection from infection conferred by probiotics. We then use the model to explore how host-mediated interventions—namely, shifts in the supply rates of electron donors (e.g., dietary fiber) and respiratory electron acceptors (e.g., oxygen)—can potentially be used to direct gut community assembly. Our study demonstrates how resource competition and ecological feedbacks between the host and the gut microbiota can be critical determinants of human health outcomes. We identify several testable model predictions ready for experimental validation. [ABSTRACT FROM AUTHOR]

Copyright of American Naturalist is the property of University of Chicago and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Plants present a variety of defensive strategies against herbivores, broadly classified into tolerance and resistance. Since resource availability can also limit plant growth, we expect plant allocation to resource acquisition and defense to vary along resource gradients. Yet, the conditions under which one defensive strategy is favored over the other are unclear. Here, we use an eco-evolutionary model to investigate plant adaptive allocation to resource acquisition, tolerance, and resistance along a resource gradient in a simple food web module inspired by plankton communities where plants compete for a single resource and are grazed on by a shared herbivore. We show that undefended, acquisition-specialist strategies dominate under low resource supplies. Conversely, high resource supplies, which lead to high herbivore abundance because of trophic transfers, result in either the dominance of very resistant strategies or coexistence between a completely resistant strategy and a fast-growing, tolerant one. We also explore the consequences of this adaptive allocation on species biomasses. Finally, we compare our predictions to a more traditional, density-independent optimization model. We show that density dependence mediated by resources and herbivores is the cause of the increase in plant resistance along the resource gradient, as the optimization model would instead have favored tolerance. [ABSTRACT FROM AUTHOR]

To persist in a changing world, populations must adapt. The ability to adapt is influenced by interactions with other species, such as predators. Recent experiments and theory suggest that selective pressures arising from predation may help prey adapt phenotypically to changing environments, but how this influences persistence remains unclear. In particular, it has not yet been shown whether predator-induced adaptation can outweigh predator-imposed reductions in population size, allowing prey to persist when they would otherwise go extinct. Here we examine if (and if so, how) predation can enhance the ability of prey to persist in a directionally changing environment. To do so, we extend a single-species quantitative-genetics framework that predicts rates of environmental change beyond which populations go extinct. While we assume predation decreases prey density, we find that predators can indeed help prey persist if they sufficiently increase prey adaptedness (decrease phenotypic lag). We show two ways this can occur: (1) the selective push, in which predators consume maladapted individuals and thus add selection that pushes the mean prey trait toward its optimum; and (2) the evolutionary hydra effect, when predation reduces prey density and thereby increases prey birthrate, allowing more selective events per unit time and effectively reducing generation time. We also discuss how our results apply more broadly to sources of mortality beyond predation. [ABSTRACT FROM AUTHOR]

Spatial structure can decisively influence the way evolutionary processes unfold. To date, several methods have been used to study evolution in spatial systems, including population genetics, quantitative genetics, moment-closure approximations, and individual-based models. Here we extend the study of spatial evolutionary dynamics to eco-evolutionary models based on reaction-diffusion equations and adaptive dynamics. Specifically, we derive expressions for the strength of directional and stabilizing/disruptive selection that apply both in continuous space and to metacommunities with symmetrical dispersal between patches. For directional selection on a quantitative trait, this yields a way to integrate local directional selection across space and determine whether the trait value will increase or decrease. The robustness of this prediction is validated against quantitative genetics. For stabilizing/disruptive selection, we show that spatial heterogeneity always contributes to disruptive selection and hence always promotes evolutionary branching. The expression for directional selection is numerically very efficient and hence lends itself to simulation studies of evolutionary community assembly. We illustrate the application and utility of the expressions for this purpose with two examples of the evolution of resource utilization. Finally, we outline the domain of applicability of reaction-diffusion equations as a modeling framework and discuss their limitations. [ABSTRACT FROM AUTHOR]

Most mutualisms in nature involve interactions between multispecies mutualist guilds and multiple partner species. While mechanisms such as niche partitioning can explain part of this diversity, the presence of low-quality partners, which produce relatively low returns on investment compared with other guild members, is not well understood. Here, we consider a novel explanation for this persistence: that low-quality partners are actively maintained by their hosts as part of a growth-maximizing strategy, even in the presence of higher-quality alternatives. We use a model inspired by the interaction between host trees and ectomycorrhizal fungi to demonstrate that when the environment is variable, trees maintain low-quality fungal partners that they would not otherwise maintain in constant environments. This active investment, which emerges as a response to saturating returns on investment in higher-quality partners, could contribute to the maintenance of diversity in multi-species mutualisms. [ABSTRACT FROM AUTHOR]

Anthropogenic environmental change can affect species directly by altering physiological rates or indirectly by changing competitive outcomes. The unknown strength of competition-mediated indirect effects makes it difficult to predict species abundances in the face of ongoing environmental change. Theory developed with phenomenological competition models shows that indirect effects are weak when coexistence is strongly stabilized, but these models lack a mechanistic link between environmental change and species performance. To extend existing theory, we examined the relationship between coexistence and indirect effects in mechanistic resource competition models. We defined environmental change as a change in resource supply points and quantified the resulting competition-mediated indirect effects on species abundances. We found that the magnitude of indirect effects increases in proportion to niche overlap. However, indirect effects also depend on differences in how competitors respond to the change in resource supply, an insight hidden in nonmechanistic models. Our analysis demonstrates the value of using niche overlap to predict the strength of indirect effects and clarifies the types of indirect effects that global change can have on competing species. [ABSTRACT FROM AUTHOR]

Recent studies investigating feedbacks between evolution and ecology suggest that microevolution may affect community structure.Motivated by this, we use a quantitative genetics and Lotka-Volterra framework to understand the impact of eco-evolutionary feedbacks on an intraguild predation community in which the intraguild predator evolves between a phenotype specialized for attacking its competitor (the intraguild prey) and a phenotype specialized for attacking a common resource. We show that evolution can drive both sudden and gradual shifts in community structure. Evolutionary rescue of the community, in which evolution prevents the loss of species, occurs in two ways: (i) selection to a particular phenotype that supports coexistence and (ii) continuous evolution between prey- and resource-specialist phenotypes that support communitylevel Red Queen dynamics in which the community fluctuates between prey- and predator-dominated states. Paradoxically, the predator can evolve to extirpate itself from the community provided that the ecological dynamics support bistability. Strong trade-offs between specialist phenotypes can lead to trait-based alternative states of the community resembling either a food chain or exploitative competition. Finally, we show that rapid evolution can stabilize equilibria that are unstable for the ecological dynamics, whereas slow evolution can stabilize equilibria that are unstable for the evolutionary dynamics. Our results demonstrate that eco-evolutionary feedbacks can drive shifts in community structure and that the overall dynamics depend on the trade-off strength and evolutionary rate. [ABSTRACT FROM AUTHOR]

The resources that organisms depend on often fluctuate over time, and a variety of common traits are thought to be adaptations to variable resource supply. To understand the trait structure of communities, it is necessary to understand the functional tradeoffs that determine what trait combinations are possible and which species can persist and coexist in a given environment. We compare traits across phytoplankton species in order to test for proposed trade-offs between maximum growth rate, equilibrium competitive ability for phosphorus (P), and ability to store P. We find evidence for a three-way trade-off between these traits, and we use empirical trait covariation to parameterize a mechanistic model of competition under pulsed P supply. The model shows that different strategies are favored under different conditions of nutrient supply regime, productivity, and mortality. Furthermore, multiple strategies typically coexist, and the range of traits that persist in the model is similar to the range of traits found in real species. These results suggest that mechanistic models informed by empirical trait variation, in combination with data on the trait structure of natural communities, will play an important role in uncovering the mechanisms that underlie the diversity and structure of ecological communities. [ABSTRACT FROM AUTHOR]

Plankton seasonal succession is a classic example of nonequilibrium community dynamics. Despite the fact that it has been well studied empirically, it lacks a general quantitative theory. Here we investigate a food web model that includes a resource, two phytoplankton, and a shared grazer-the diamond food web-in a seasonal environment. The model produces a number of successional trajectories that have been widely discussed in the context of the verbal Plankton Ecology Group model of succession, such as a spring bloom of a good competitor followed by a grazer-induced clearwater phase, setting the stage for the late-season dominance of a grazer-resistant species. It also predicts a novel, counterintuitive trajectory where the grazer-resistant species has both early- and lateseason blooms. The model often generates regular annual cycles but sometimes produces multiyear cycles or chaos, even with identical forcing each year. Parameterizing the model, we show how the successional trajectory depends on nutrient supply and the length of the growing season, two key parameters that vary among water bodies. This model extends nonequilibrium theory to food webs and is a first step toward a quantitative theory of plankton seasonal succession. [ABSTRACT FROM AUTHOR]

By definition, mutualisms involve the exchange of goods or services between partners. It has been shown that mutualism can grade into parasitism, but even when exchange is mutually beneficial, a conflict of interest remains because each partner benefits from reaping more benefits at a lower cost. Metaphorically, the partners negotiate the conditions of trade, the outcome of which will determine the net benefit to each partner. Each partner can adjust its allocation to self-provisioning while negotiating the ratio at which benefits are exchanged. To understand how these two features of trade affect mutualisms, we used the example of the plant-arbuscular mycorrhizal mutualism and modeled uptake and trade of two resources, phosphorus and carbon. In most contexts, the fungus specialized on phosphorus uptake while the plant took up both phosphorus and carbon. However, when phosphorus was abundant and light was scarce, the plant specialized, taking up only carbon and relying on trade for phosphorus. Resource availability was the most important factor determining specialization and the outcome of negotiation and trade, but other aspects of the context were also important. These results suggest experiments to link these two key features of trade with environmental conditions to determine the outcome of mutualism. [ABSTRACT FROM AUTHOR]

Phytoplankton compete for nutrients and light in a vertically heterogeneous environment determined by turbulent mixing. We analyzed a model of competition between two phytoplankton species in a stratified water column. We assume that the surface layer is uniformly mixed and that the deep layer is poorly mixed, as is commonly observed in lakes and oceans. We employed two analytical techniques, Iout- R theory in the mixed surface layer and a game theoretical approach in the deep layer. Under our assumptions, at equilibrium, each species is either absent or resides in the benthic layer, the deep layer, or the surface layer. Assuming a trade-off between nutrient- and light-competitive abilities, we obtained five spatial configurations of coexistence and the corresponding parameter regions where they occur. Good light competitors show two distinct ecological niches: in mesotrophic conditions, they live as understory species below a layer of good nutrient competitors; in eutrophic conditions, they live as competitive dominants in the surface layer. Multiple regions of alternative stable states are possible in parameter space. This work extends previous phytoplankton competition theory to stratified water columns, as commonly found in lakes and oceans. [ABSTRACT FROM AUTHOR]

Planktonic microorganisms are affected by various size-dependent processes both from the bottom up and from the top down. We developed a simple resource-consumer model to explore how size-dependent resource uptake and resource loss influence the growth of, and competition between, planktonic microorganisms. We considered three steps of resource uptake: diffusive transport of resource molecules, uptake by membrane transporters, and cellular enzymatic catalysis, and we investigated optimal cell size when one, two, or three of those steps limit resource uptake. Optimal cell size depends negatively on the size of resource molecules when resource uptake is limited by diffusive transport and membrane uptake. When competing for two resources of different molecular sizes, two different-sized consumers can coexist if the inputs of resources and sizes of consumers are correctly chosen. The model suggests that mixtures of various-sized resources can promote coexistence and size diversity of microorganisms even if the availability of one element, such as carbon, nitrogen, or phosphorus, limits the whole community. Model predictions include that bacteria grown on maltose or polysaccharides should be smaller compared with those grown on glucose under carbon limitation. Our results suggest that size of resource molecules can be an important factor in microbial resource competition in aquatic environments. [ABSTRACT FROM AUTHOR]

Recent theoretical studies have shown that spatial redistribution of surface water may explain the occurrence of patterns of alternating vegetated and degraded patches in semiarid grasslands. These results implied, however, that spatial redistribution processes cannot explain the collapse of production on coarser scales observed in these systems. We present a spatially explicit vegetation model to investigate possible mechanisms explaining irreversible vegetation collapse on coarse spatial scales. The model results indicate that the dynamics of vegetation on coarse scales are determined by the interaction of two spatial feedback processes. Loss of plant cover in a certain area results in increased availability of water in remaining vegetated patches through run-on of surface water, promoting within-patch plant production. Hence, spatial redistribution of surface water creates negative feedback between reduced plant cover and increased plant growth in remaining vegetation. Reduced plant cover, however, results in focusing of herbivore grazing in the remaining vegetation. Hence, redistribution of herbivores creates positive feedback between reduced plant cover and increased losses due to grazing in remaining vegetated patches, leading to collapse of the entire vegetation. This may explain irreversible vegetation shifts in semiarid grasslands on coarse spatial scales. [ABSTRACT FROM AUTHOR]

Evaluates how light fluctuations affect phytoplankton community structure and diversity. Different types of light fluctuations; Effect of seasonal change in day length on the seasonal succession of phytoplankton; Coexistence regions for phytoplankton species.

Presents a study that investigates two assumptions of models dealing with the loss of habitat due to human activities. Reference to D. Tilman et al.'s (1994, 1997) work on the colonization rates of successive species in the competitive hierarchy; Failure of Tilman's model to account for the spatial location of sites on the landscapes; Methodology used in the investigations.