The noisy edge of traveling waves

Traveling waves are ubiquitous in nature and control the speed of many important dynamical processes, including chemical reactions, epidemic outbreaks, and biological evolution. Despite their fundamental role in complex systems, traveling waves remain elusive because they are often dominated by rare fluctuations in the wave tip, which have defied any rigorous analysis so far. Here, we show that by adjusting nonlinear model details, noisy traveling waves can be solved exactly. The moment equations of these tuned models are closed and have a simple analytical structure resembling the deterministic approximation supplemented by a nonlocal cutoff term. The peculiar form of the cutoff shapes the noisy edge of traveling waves and is critical for the correct prediction of the wave speed and its fluctuations. Our approach is illustrated and benchmarked using the example of fitness waves arising in simple models of microbial evolution, which are highly sensitive to number fluctuations. We demonstrate explicitly how these models can be tuned to account for finite population sizes and determine how quickly populations adapt as a function of population size and mutation rates. More generally, our method is shown to apply to a broad class of models, in which number fluctuations are generated by branching processes. Because of this versatility, the method of model tuning may serve as a promising route toward unraveling universal properties of complex discrete particle systems.Comment: For supplementary material and published open access article, see http://www.pnas.org/content/108/5/1783.abstract?sid=693e63f3-fd1a-407a-983e-c521efc6c8c5 See also Commentary Article by D. S. Fisher, http://www.pnas.org/content/108/7/2633.extrac

Life at the front of an expanding population

Recent microbial experiments suggest that enhanced genetic drift at the frontier of a two-dimensional range expansion can cause genetic sectoring patterns with fractal domain boundaries. Here, we propose and analyze a simple model of asexual biological evolution at expanding frontiers to explain these neutral patterns and predict the effect of natural selection. Our model attributes the observed gradual decrease in the number of sectors at the leading edge to an unbiased random walk of sector boundaries. Natural selection introduces a deterministic bias in the wandering of domain boundaries that renders beneficial mutations more likely to escape genetic drift and become established in a sector. We find that the opening angle of those sectors and the rate at which they become established depend sensitively on the selective advantage of the mutants. Deleterious mutations, on the other hand, are not able to establish a sector permanently. They can, however, temporarily "surf" on the population front, and thereby reach unusual high frequencies. As a consequence, expanding frontiers are susceptible to deleterious mutations as revealed by the high fraction of mutants at mutation-selection balance. Numerically, we also determine the condition at which the wild type is lost in favor of deleterious mutants (genetic meltdown) at a growing front. Our prediction for this error threshold differs qualitatively from existing well-mixed theories, and sets tight constraints on sustainable mutation rates for populations that undergo frequent range expansions.Comment: Updat

Noise Driven Evolutionary Waves

Adaptation in spatially extended populations entails the propagation of evolutionary novelties across habitat ranges. Driven by natural selection, beneficial mutations sweep through the population in a “wave of advance”. The standard model for these traveling waves, due to R. Fisher and A. Kolmogorov, plays an important role in many scientific areas besides evolution, such as ecology, epidemiology, chemical kinetics, and recently even in particle physics. Here, we extend the Fisher–Kolmogorov model to account for mutations that confer an increase in the density of the population, for instance as a result of an improved metabolic efficiency. We show that these mutations invade by the action of random genetic drift, even if the mutations are slightly deleterious. The ensuing class of noise-driven waves are characterized by a wave speed that decreases with increasing population sizes, contrary to conventional Fisher–Kolmogorov waves. When a trade-off exists between density and growth rate, an evolutionary optimal population density can be predicted. Our simulations and analytical results show that genetic drift in conjunction with spatial structure promotes the economical use of limited resources. The simplicity of our model, which lacks any complex interactions between individuals, suggests that noise-induced pattern formation may arise in many complex biological systems including evolution

Tension dynamics in semiflexible polymers. II. Scaling solutions and applications

In part I O. Hallatschek , preceding paper, Phys. Rev. E 75, 031905 (2007)] of this contribution, a systematic coarse-grained description of the dynamics of a weakly bending semiflexible polymer was developed. Here, we discuss analytical solutions of the established deterministic partial integro-differential equation for the spatiotemporal relaxation of the backbone tension. For prototypal experimental situations, such as the sudden application or release of a strong external pulling force, it is demonstrated that the tensile dynamics reflects the self-affine conformational fluctuation spectrum in a variety of intermediate asymptotic power laws. Detailed and explicit analytical predictions for the tension propagation and relaxation and corresponding results for common observables, such as the end-to-end distance, are obtained

Coupling of transverse and longitudinal response in stiff polymers

The time-dependent transverse response of stiff polymers, represented as weakly-bending wormlike chains (WLCs), is well-understood on the linear level, where transverse degrees of freedom evolve independently from the longitudinal ones. We show that, beyond a characteristic time scale, the nonlinear coupling of transverse and longitudinal motion in an inextensible WLC significantly weakens the polymer response compared to the widely used linear response predictions. The corresponding feedback mechanism is rationalized by scaling arguments and quantified by a multiple scale approach that exploits an inherent separation of transverse and longitudinal correlation length scales. Crossover scaling laws and exact analytical and numerical solutions for characteristic response quantities are derived for different experimentally relevant setups. Our findings are applicable to cytoskeletal filaments as well as DNA under tension.Comment: 4 pages, 3 figures, 1 table; final versio