Pluralism in Evolutionary Theory

The review by Waxman and Gavrilets (Waxman and Gavrilets 2005) illustrates the collision of different mindsets in evolutionary theory. These differences originate from the awe-inspiring complexity of the evolutionary process itself: evolutionary understanding critically depends on processes at many biological levels. Starting out with base pairs and their sequences, scholars of evolution have to consider -- in the order of biological complexity -- alleles, quantitative allelic traits, physiological and morphological traits, life-history traits, demographic rates, fitness, changes in genotype frequencies, population population dynamics, trait substitution sequences, and population bifurcations, to eventually arrive at the levels of ecological communities and the biosphere. It would appear that no other field of contemporary science sports comparable ambitions

Evolutionary Branching and Sympatric Speciation Caused by Different Types of Ecological Interactions [Revised 18 September 2000]

Evolutionary branching occurs when frequency-dependent selection splits a phenotypically monomorphic population into two distinct phenotypic clusters. A prerequisite for evolutionary branching is that directional selection drives the population towards a fitness minimum in phenotype space. This paper demonstrates that selection regimes leading to evolutionary branching readily arise from a wide variety of different ecological interactions within and between species. We use classical ecological models for symmetric and asymmetric competition, for mutualism, and for predator-prey interactions to describe evolving populations with continuously varying characters. For these models, we investigate the ecological and evolutionary conditions that allow for evolutionary branching and establish that branching is a generic and robust phenomenon. Evolutionary branching becomes a model for sympatric speciation when population genetics and mating mechanisms are incorporated into ecological models. In sexual populations with random mating, the continual production of intermediate phenotypes from two incipient branches prevents evolutionary branching. In contrast, when mating is assortative for the ecological characters under study, evolutionary branching is possible in sexual populations and can lead to speciation. Therefore, we also study the evolution of assortative mating as a quantitative character. We show that evolution under branching conditions selects for assortativeness and thus allows sexual populations to escape from fitness minima. We conclude that evolutionary branching offers a general basis for understanding adaptive speciation and radiation under a wide range of different ecological conditions

A Tale of Two Cycles - Distinguishing Quasi-cycles and Limit Cycles in the Finite Predator-Prey Populations

Periodic predator-prey dynamics in constant environments are usually taken as indicative of deterministic limit cycles. It is known, however, that demographic stochasticity in finite populations can also give rise to regular population cycles, even when the corresponding deterministic models predict a stable equilibrium. Specifically, such quasi-cycles are expected in stochastic versions of deterministic models exhibiting equilibrium dynamics with weakly damped oscillations. The existence of quasi-cycles substantially expands the scope for natural patterns of periodic population oscillations caused by ecological interactions, thereby complicating the conclusive interpretation of such patterns. Here we show how to distinguish between quasi-cycles and noisy limit cycles based on observing changing population sizes in predator-prey populations. We start by confirming that both types of cycle can occur in the individual-based version of a widely used class of deterministic predator-prey model. We then show that it is feasible and straightforward to accurately distinguish between the two types of cycle through the combined analysis of autocorrelations and marginal distributions of population sizes. Finally, by confronting these results with real ecological time series, we demonstrate that by using our methods even short and imperfect time series allow quasi-cycles and limit cycles to be distinguished reliably

What We Have Also Learned: Adaptive Speciation is Theoretically Possible

A recent Perspectives article by Gavrilets (2003) on the theory of speciation ignored advances in understanding processes of adaptive speciation, in which the splitting of lineages is an adaptation caused by frequency-dependent selection. Adaptive, or sympatric, speciation has been modeled since the 1960s, but the large amount of attention from both empirical and theoretical biologists that adaptive speciation has received in recent years goes far beyond what was described in Gavrilets' paper. Due to conceptual advances based on the theory of adaptive dynamics, adaptive speciation has emerged as a theoretically plausible evolutionary process that can occur in many different ecological settings

Reputation-based conditional interaction supports cooperation in well-mixed prisoner's dilemmas

In the well-mixed prisoner's dilemma game, individuals are typically assumed to have no choice about whether to interact with other individuals in the population. In this paper, we instead consider reputation- based conditional interaction and its consequences for the evolution of cooperation. Each individual has a tolerance range, and only interacts with other individuals whose reputation lies within its tolerance range in a chosen sample of the population. Reputation contains information about the number of interaction partners an individual has just cooperated with. We find that the introduction of conditional interaction promotes cooperation in well-mixed populations, and there exist moderate tolerance ranges for which this effect is maximized. For a given tolerance range, there is a critical cost-to-benefit ratio below which cooperation can be promoted. Interestingly, we find that if cooperation evolves, different cooperators' interaction clusters are typically maintained in the population, each around a different reputation level. We further investigate some properties of these cooperators' clusters. Moreover, we examine the effects of the sample number on the evolution of cooperation. Our results highlight the importance of the detailed consideration of modes of interaction for the evolution of cooperation in well-mixed popultions

Small world effects in evolution

For asexual organisms point mutations correspond to local displacements in the genotypic space, while other genotypic rearrangements represent long-range jumps. We investigate the spreading properties of an initially homogeneous population in a flat fitness landscape, and the equilibrium properties on a smooth fitness landscape. We show that a small-world effect is present: even a small fraction of quenched long-range jumps makes the results indistinguishable from those obtained by assuming all mutations equiprobable. Moreover, we find that the equilibrium distribution is a Boltzmann one, in which the fitness plays the role of an energy, and mutations that of a temperature.Comment: 13 pages and 5 figures. New revised versio

How Gaussian competition leads to lumpy or uniform species distributions

A central model in theoretical ecology considers the competition of a range of species for a broad spectrum of resources. Recent studies have shown that essentially two different outcomes are possible. Either the species surviving competition are more or less uniformly distributed over the resource spectrum, or their distribution is 'lumped' (or 'clumped'), consisting of clusters of species with similar resource use that are separated by gaps in resource space. Which of these outcomes will occur crucially depends on the competition kernel, which reflects the shape of the resource utilization pattern of the competing species. Most models considered in the literature assume a Gaussian competition kernel. This is unfortunate, since predictions based on such a Gaussian assumption are not robust. In fact, Gaussian kernels are a border case scenario, and slight deviations from this function can lead to either uniform or lumped species distributions. Here we illustrate the non-robustness of the Gaussian assumption by simulating different implementations of the standard competition model with constant carrying capacity. In this scenario, lumped species distributions can come about by secondary ecological or evolutionary mechanisms or by details of the numerical implementation of the model. We analyze the origin of this sensitivity and discuss it in the context of recent applications of the model.Comment: 11 pages, 3 figures, revised versio

On the evolution of decoys in plant immune systems

The Guard-Guardee model for plant immunity describes how resistance proteins (guards) in host cells monitor host target proteins (guardees) that are manipulated by pathogen effector proteins. A recently suggested extension of this model includes decoys, which are duplicated copies of guardee proteins, and which have the sole function to attract the effector and, when modified by the effector, trigger the plant immune response. Here we present a proof-of-principle model for the functioning of decoys in plant immunity, quantitatively developing this experimentally-derived concept. Our model links the basic cellular chemistry to the outcomes of pathogen infection and resulting fitness costs for the host. In particular, the model allows identification of conditions under which it is optimal for decoys to act as triggers for the plant immune response, and of conditions under which it is optimal for decoys to act as sinks that bind the pathogen effectors but do not trigger an immune response.Comment: 15 pages, 6 figure

The influence of habitat boundaries on evolutionary branching along environmental gradients

It is well known that habitat boundaries affect ecological dynamics, but their influence on evolutionary dynamics is less well understood. Here, we study the effects of different kinds of boundaries on evolutionary branching in clonal populations along environmental gradients by systematically analyzing individual-based stochastic models in small- and large-range systems, as well as their large-population-size limits through deterministic approximations. Specifically, we examine four prototypical kinds of boundaries: impermeable boundaries at which individuals stop (“stopping”), or from which they continue back into the interior as if bouncing back mechanically (“reflecting”), or that let them abort the dispersal attempt, return to their original position and try a different direction (“reprising”), and semipermeable boundaries that can be crossed without hindrance, but do not allow the crossing individual to return (“absorbing”).We find that boundary conditions shape branching patterns only in small-range systems, where stopping boundaries generate disruptive selection for a wide range of parameters, whereas absorbing boundaries always generate stabilizing selection. Reflecting and reprising boundaries generate disruptive selection at low individual mobilities, and stabilizing selection at high mobilities. To further analyze these findings, we introduce a simple approximation of the invasion fitness in a mobile population, which predicts the observed outcome. The effect of stochasticity on evolutionary outcomes is small even in small populations: stochasticity causes random branch extinctions at steeper slopes and higher mobilities. In large-range systems, frequency-dependent interactions alone induce evolutionary branching for almost all parameters and independent of boundary conditions