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Lehmann, L.

Publications and source records attributed to Lehmann, L..

45 records · Page 3Linked to original sources

Invasion implies substitution in ecological communities with class-structured populations

Long-term evolution of quantitative traits is classically and usefully described as the directional change in phenotype due to the recurrent fixation of new mutations. A formal justification for such continual evolution ultimately relies on the "invasion implies substitution"-principle. Here, whenever a mutant allele causing a small phenotypic change can successfully invade a population, the ancestral (or wild-type) allele will be replaced, whereby fostering gradual phenotypic change if the process is repeated. It has been argued that this principle holds in a broad range of situations, including spatially and demographically structured populations experiencing frequency and density dependent selection under demographic and environmental fluctuations. However, prior studies have not been able to account for all aspects of population structure, leaving unsettled the conditions under which the "invasion implies substitution"-principle really holds. In this paper, we start by laying out a program to explore and clarify the generality of the "invasion implies substitution"-principle. Particular focus is given on finding an explicit and functionally constant representation of the selection gradient on a quantitative trait. Using geometric singular perturbation methods, we then show that the "invasion implies substitution"-principle generalizes to well-mixed and scalar-valued polymorphic multispecies ecological communities that are structured into finitely many demographic (or physiological) classes. The selection gradient is shown to be constant over the evolutionary timescale and that it depends only on the resident phenotype, individual growth-rates, population steady states and reproductive values, all of which are calculated from the resident dynamics. Our work contributes to the theoretical foundations of evolutionary ecology.

evolutionary biology↗

When do individuals maximize their inclusive fitness?

Adaptation is often described in behavioral ecology as individuals maximizing their inclusive fitness. Under what conditions does this hold and how does this relate to the gene-centered perspective of adaptation? We unify and extend the literature on these questions to class-structured populations. We demonstrate that the maximization (in the best-response sense) of class-specific inclusive fitness obtains in uninvadable population states (meaning that all deviating mutant go extinct). This defines a genuine actor-centered perspective on adaptation. But this inclusive fitness is assigned to all bearers of a mutant allele in a given class and depends on distributions of demographic and genetic contexts. These distributions, in turn, usually depend on events in previous generations and are thus not under individual control. This prevents, in general, from envisioning individuals themselves as autonomous fitness-maximizers, each with its own inclusive fitness. For weak selection, however, the dependence on earlier events can be neglected. We then show that each individual in each class appears to maximize its own inclusive fitness when all other individuals exhibit fitness-maximizing behavior. This defines a genuine individual-centered perspective of adaptation and justifies formally, as a first-order approximation, the long-heralded view of individuals appearing to maximize their own inclusive fitness.

evolutionary biology↗

Linking micro and macroevolution in the presence of migration

1. The process of speciation is of key importance in evolutionary biology because it shapes macroevolutionary patterns. This process starts at the microevolutionary level, for instance, when two subpopulations evolve towards different phenotypic optima. The speed at which these optima are reached is controlled by the degree of stabilising selection, which pushes a mean trait towards an optimum within subpopulations, and ongoing migration that pulls the mean phenotype away from that optimum. Traditionally, macro phenotypic evolution with selection has been modelled by Ornstein-Uhlenbeck (OU) processes, but these models have ignored the role of migration within species.\n\n2. Here, our goal is to reconcile the processes of micro and macroevolution by modelling migration during speciation. More precisely, we introduce an OU model where migration happens between two subpopulations within a branch of a phylogeny and this migration decreases over time as it happens during speciation. We then use this model to study the evolution of trait means along a phylogeny, as well as the way phenotypic disparity between species changes with successive epochs.\n\n3. We show that ignoring the effect of migration in sampled time-series data leads to a significant underestimation of the selective forces acting upon it. We also show that migration decreases the expected phenotypic disparity between species and we show the effect of migration in the particular case of niche filling. We further introduce a method to jointly estimate selection and migration from time-series data.\n\n4. Our model extends standard results of interactions selection-migration in a microevolutionary time frame across multiple speciation events at a macroevolutionary scale. Our results further proof that not accounting for gene flow has important consequences in inferences at both the micro and macroevolutionary scale.

evolutionary biology↗

Sex allocation conflict and sexual selection throughout the lifespan of eusocial colonies

Models of sex allocation conflict are central to evolutionary biology but have mostly assumed static decisions, where resource allocation strategies are constant over colony lifespan. Here, we develop a model to study how the evolution of dynamic resource allocation strategies is affected by the queen-worker conflict in annual eusocial insects. We demonstrate that the time of dispersal of sexuals affects the sex allocation ratio through sexual selection on males. Furthermore, our model provides three predictions that depart from established results of classic static allocation models. First, we find that the queen wins the sex allocation conflict, while the workers determine the maximum colony size and colony productivity. Second, male-biased sex allocation and protandry evolve if sexuals disperse directly after eclosion. Third, when workers are more related to new queens, then the proportional investment into queens is expected to be lower, which results from the interacting effect of sexual selection (selecting for protandry) and sex allocation conflict (selecting for earlier switch to producing sexuals). Overall, we find that colony ontogeny crucially affects the outcome of sex-allocation conflict because of the evolution of distinct colony growth phases, which decouples how queens and workers affect allocation decisions and can result in asymmetric control.

evolutionary biology↗

Evolution of preferences in group-structured populations: genes, guns, and culture

During human evolution, individuals interacted mostly within small groups that were connected by limited migration and sometimes by conflicts. Which preferences, if any, will prevail in such scenarios? Building on population biology models of spatially structured populations, and assuming individuals preferences to be their private information, we characterize those preferences that, once established, cannot be displaced by alternative preferences. We represent such uninvadable preferences in terms of fitness and in terms of material payoffs. At the fitness level, individuals can be regarded to act as if driven by a mix of self-interest and a Kantian motive that evaluates own behavior in the light of the consequences for own fitness if others adopted this behavior. This Kantian motive is borne out from (genetic or cultural) kin selection. At the material-payoff level, individuals act as if driven in part by self-interest and a Kantian motive (in terms of material payoffs), but also in part by other-regarding preferences towards other group members. This latter motive is borne out of group resource constraints and the risk of conflict with other groups. We show how group size, the migration rate, the risk of group conflicts, and cultural loyalty shape the relative strengths of these motives.\n\nJEL codesA12, A13, B52, C73, D01, D63, D64, D91.

evolutionary biology↗

A quantitative genetics model for the dynamics of phenotypic (co)variances under limited dispersal, with an application to the coevolution of socially synergistic traits

Darwinian evolution consists of the gradual transformation of heritable quantitative traits due to natural selection and the input of random variation by mutation. Here, we use a quantitative genetics approach to investigate the coevolution of multiple traits under selection, mutation, and limited dispersal. We track the dynamics of trait means and variance-covariances between traits that experience frequency-dependent selection. Assuming a multivariate-normal trait distribution, we recover classical dynamics of quantitative genetics, as well as stability and evolutionary branching conditions of invasion analyses, except that due to limited dispersal, selection depends on indirect fitness effects and relatedness. In particular, correlational selection that associates different traits within-individuals depends on the fitness effects of such associations between-individuals. These kin selection effects can be as relevant as pleiotropy for correlation between traits. We illustrate this with an example of the coevolution of two social traits whose association within-individual is costly but synergistically beneficial between-individuals. As dispersal becomes limited and relatedness increases, associations between-traits between-individuals become increasingly targeted by correlational selection. Consequently, the trait distribution goes from being bimodal with a negative correlation under panmixia to unimodal with a positive correlation under limited dispersal. More broadly, our approach can help understand the evolution of intra-specific variation.

evolutionary biology↗

Eco-evolutionary dynamics under limited dispersal: ecological inheritance, altruism within and spite between species

Understanding selection on ecological interactions that take place in dispersal-limited communities is an important challenge for ecology and evolutionary biology. The problem is that local demographic stochasticity generates eco-evolutionary dynamics that are too complicated to make tractable analytical investigations. Here, we circumvent this problem by approximating the selection gradient on a quantitative trait that influences local community dynamics, assuming that such dynamics are deterministic with a stable fixed point, but incorporating the kin selection effects arising from demographic stochasticity. Our approximation reveals that selection depends on how an individual expressing a trait-change influences: (1) its own fitness and the fitness of its current relatives; and (2) the fitness of its downstream relatives through modifications of local ecological conditions (i.e., through ecological inheritance). Mathematically, the effects of ecological inheritance on selection are captured by dispersal-limited versions of press-perturbations of community ecology. We use our approximation to investigate the evolution of helping within- and harming between-species when these behaviours influence demography. We find helping evolves more readily when competition is for material resources rather than space because then, the costs of kin competition are paid by downstream relatives. Similarly, harming between species evolves when it alleviates downstream relatives from inter-specific competition. Beyond these examples, our approximation can help better understand the influence of ecological inheritance on a variety of eco-evolutionary dynamics, from plant-pollinator to predator-prey coevolution.

evolutionary biology↗

Co-evolution of dispersal with behaviour favours social polymorphism

Dispersal determines gene flow among groups in a population and so plays a major role in many ecological and evolutionary processes, from biological invasions to species extinctions. Because patterns of gene flow shape kin structure, dispersal is also important to the evolution of social behaviours that influence reproduction and survival within groups. Conversely, dispersal patterns depend on kin structure and social behaviour. Dispersal and social behaviour therefore co-evolve but the nature and consequences of this interplay are not well understood. Here, we model this co-evolution and show that it readily leads to the emergence and maintenance of two broadly-defined social morphs: a sessile, benevolent morph expressed by individuals who tend to increase the fecundity of others within their group relative to their own; and a dispersive, selfserving morph expressed by individuals who tend to increase their own fecundity relative to others within their group. This social polymorphism arises as a consequence of a positive linkage between the loci responsible for dispersal and social behaviour, leading to benevolent individuals preferentially interacting with relatives and self-serving individuals with non-relatives. We find that this positive linkage is favoured under a large spectrum of conditions, which suggests that an association between dispersal proclivity and other social traits should be common in nature. In line with this prediction, dispersing individuals across a wide range of organisms have been reported to differ in their social tendencies from non-dispersing individuals.

evolutionary biology↗

Invasion fitness for gene-culture co-evolution in the family and an application to cumulative culture under vertical transmission

Human evolution depends on the co-evolution between genetically determined behaviors and socially transmitted information. Although vertical transmission of cultural information from parent to off-spring is common in hominins, its effects on cumulative cultural evolution are not fully understood. Here, we investigate gene-culture co-evolution in a family-structured population by studying the invasion fitness of a mutant allele that influences a deterministic level of cultural information (e.g., amount of knowledge or skill) to which diploid carriers of the mutant are exposed in subsequent generations. We show that the selection gradient on such a mutant, and the concomitant level of cultural information it generates, can be evaluated analytically under the assumption that the cultural dynamic has a single attractor point, thereby making gene-culture co-evolution in family-structured populations with multigenerational effects mathematically tractable. We apply our result to study how genetically determined phenotypes of individual and social learning co-evolve with the level of adaptive information they generate under vertical transmission. We find that vertical transmission increases adaptive information due to kin selection effects, but when information is transmitted as efficiently between family members as between unrelated individuals, this increase is moderate in diploids. By contrast, we show that the way resource allocation into learning trades off with allocation into reproduction (the \"learning-reproduction trade-off\") significantly influences levels of adaptive information. We also show that vertical transmission prevents evolutionary branching and may therefore play a qualitative role in gene-culture co-evolutionary dynamics. More generally, our analysis of selection suggests that vertical transmission can significantly increase levels of adaptive information under the biologically plausible condition that information transmission between relatives is more efficient than between unrelated individuals.

evolutionary biology↗