Universality of fixation probabilities in randomly structured populations

The stage of evolution is the population of reproducing individuals. The structure of the population is known to affect the dynamics and outcome of evolutionary processes, but analytical results for generic random structures have been lacking. The most general result so far, the isothermal theorem, assumes the propensity for change in each position is exactly the same, but realistic biological structures are always subject to variation and noise. We consider a finite population under constant selection whose structure is given by a variety of weighted, directed, random graphs; vertices represent individuals and edges interactions between individuals. By establishing a robustness result for the isothermal theorem and using large deviation estimates to understand the typical structure of random graphs, we prove that for a generalization of the Erdős-Rényi model, the fixation probability of an invading mutant is approximately the same as that of a mutant of equal fitness in a well-mixed population with high probability. Simulations of perturbed lattices, small-world networks, and scale-free networks behave similarly. We conjecture that the fixation probability in a well-mixed population, (1 − r−1)/(1 − r−n), is universal: for many random graph models, the fixation probability approaches the above function uniformly as the graphs become large

Amplifiers of selection

When a new mutant arises in a population, there is a probability it outcompetes the residents and fixes. The structure of the population can affect this fixation probability. Suppressing population structures reduce the difference between two competing variants, while amplifying population structures enhance the difference. Suppressors are ubiquitous and easy to construct, but amplifiers for the large population limit are more elusive and only a few examples have been discovered. Whether or not a population structure is an amplifier of selection depends on the probability distribution for the placement of the invading mutant. First, we prove that there exist only bounded amplifiers for adversarial placement-that is, for arbitrary initial conditions. Next, we show that the Star population structure, which is known to amplify for mutants placed uniformly at random, does not amplify for mutants that arise through reproduction and are therefore placed proportional to the temperatures of the vertices. Finally, we construct population structures that amplify for all mutational events that arise through reproduction, uniformly at random, or through some combination of the two

The Time Scale of Evolutionary Innovation

A fundamental question in biology is the following: what is the time scale that is needed for evolutionary innovations? There are many results that characterize single steps in terms of the fixation time of new mutants arising in populations of certain size and structure. But here we ask a different question, which is concerned with the much longer time scale of evolutionary trajectories: how long does it take for a population exploring a fitness landscape to find target sequences that encode new biological functions? Our key variable is the length, of the genetic sequence that undergoes adaptation. In computer science there is a crucial distinction between problems that require algorithms which take polynomial or exponential time. The latter are considered to be intractable. Here we develop a theoretical approach that allows us to estimate the time of evolution as function of We show that adaptation on many fitness landscapes takes time that is exponential in even if there are broad selection gradients and many targets uniformly distributed in sequence space. These negative results lead us to search for specific mechanisms that allow evolution to work on polynomial time scales. We study a regeneration process and show that it enables evolution to work in polynomial time

Cellular cooperation with shift updating and repulsion

Population structure can facilitate evolution of cooperation. In a structured population, cooperators can form clusters which resist exploitation by defectors. Recently, it was observed that a shift update rule is an extremely strong amplifier of cooperation in a one dimensional spatial model. For the shift update rule, an individual is chosen for reproduction proportional to fecundity; the offspring is placed next to the parent; a random individual dies. Subsequently, the population is rearranged (shifted) until all individual cells are again evenly spaced out. For large population size and a one dimensional population structure, the shift update rule favors cooperation for any benefit-to-cost ratio greater than one. But every attempt to generalize shift updating to higher dimensions while maintaining its strong effect has failed. The reason is that in two dimensions the clusters are fragmented by the movements caused by rearranging the cells. Here we introduce the natural phenomenon of a repulsive force between cells of different types. After a birth and death event, the cells are being rearranged minimizing the overall energy expenditure. If the repulsive force is sufficiently high, shift becomes a strong promoter of cooperation in two dimensions