Search

Many social interactions happen indirectly via modifications of the environment, e.g. through the secretion of functional compounds or the depletion of renewable resources. Here, we derive the selection gradient on a quantitative trait affecting dynamical environmental variables that feed back on reproduction and survival in a finite patch-structured population subject to isolation by distance. Our analysis shows that the selection gradient depends on how a focal individual influences the fitness of all future individuals in the population through modifications of the environmental variables they experience, weighted by the neutral relatedness between recipients and the focal. The evolutionarily relevant trait-driven environmental modifications are formalized as the extended phenotypic effects of an individual, quantifying how a trait change in an individual in the present affects the environmental variables in all patches at all future times. When the trait affects reproduction and survival through a payoff function, the selection gradient can be expressed in terms of extended phenotypic effects weighted by scaled relatedness. We show how to compute extended phenotypic effects, relatedness, and scaled relatedness using Fourier analysis, which allow us to investigate a broad class of environmentally mediated social interactions in a tractable way. We use our approach to study the evolution of a trait controlling the costly production of some lasting commons (e.g. a common-pool resource or a toxic compound) that can diffuse in space and persist in time. We show that indiscriminate posthumous spite readily evolves in this scenario. More generally, whether selection favours environmentally mediated altruism or spite is determined by the spatial correlation between an individual's lineage and the commons originating from its patch. The sign of this correlation depends on interactions between dispersal patterns and the commons' renewal dynamics. More broadly, we suggest that selection can favour a wide range of social behaviours when these have carry-over effects in space and time. Author summary: Organisms continually modify their environment by extracting resources, releasing toxins, or engineering habitats. These environmental modifications can have significant fitness consequences on other organisms, including those living far away in space and the distant future. To better understand the evolutionary relevance of such environmentally mediated social interactions, we quantify natural selection on a trait that influences biotic or abiotic environmental variables in a population composed of finite patches experiencing isolation by distance, as is typical of taxa facing limited dispersal. Using Fourier analysis, we show how selection on a trait due to its environmental effects can be understood in terms of the way that a focal individual influences the fitness of all future individuals at all spatial positions via a modification to the environment these individuals encounter, whose importance depends on the relatedness between these and the focal. Simply put, natural selection tends to favour traits whose environmental effects help (resp. harm) individuals that are more (resp. less) related than average, including those distantly in space and time. In addition to allowing for a full quantification of directional selection, this inclusive fitness perspective helps garner intuition for the evolution of environmentally mediated social interactions in general and of spite in particular. [ABSTRACT FROM AUTHOR]

Biological adaptation is the outcome of allele-frequency change by natural selection. At the same time, populations are usually class structured as individuals occupy different states, such as age, sex, or stage. This is known to result in the differential transmission of alleles through nonheritable fitness differences called class transmission, which also affects allele-frequency change even in the absence of selection. How does one then isolate allele-frequency change due to selection from that due to class transmission? We decompose one-generational allele-frequency change in terms of effects of selection and class transmission and show how reproductive values can be used to reach a decomposition between any two distant generations of the evolutionary process. This provides a missing relationship between multigenerational allele-frequency change and the operation of selection. It also allows a measure of fitness to be defined summarizing the effect of selection in a multigenerational evolutionary process, which connects asymptotically to invasion fitness. [ABSTRACT FROM AUTHOR]

In patch- or habitat-structured populations, different processes can favor adaptive polymorphism at different scales. While spatial heterogeneity can generate spatially disruptive selection favoring variation between patches, local competition can lead to locally disruptive selection promoting variation within patches. So far, almost all theory has studied these two processes in isolation. Here, we use mathematical modeling to investigate how resource variation within and between habitats influences the evolution of variation in a consumer population where individuals compete in finite patches connected by dispersal. We find that locally and spatially disruptive selection typically act in concert, favoring polymorphism under a wider range of conditions than when in isolation. But when patches are small and dispersal between them is low, kin competition inhibits the emergence of polymorphism, especially when the latter is driven by local competition for resources. We further use our model to clarify what comparisons between trait and neutral genetic differentiation ( Q ST / F ST comparisons) can tell about the nature of selection. Overall, our results help us understand the interaction between two major drivers of polymorphism: locally and spatially disruptive selection, and how this interaction is modulated by the unavoidable effects of kin selection under limited dispersal. [ABSTRACT FROM AUTHOR]