Transport control by coherent zonal flows in the core/edge transitional regime

3D Braginskii turbulence simulations show that the energy flux in the core/edge transition region of a tokamak is strongly modulated - locally and on average - by radially propagating, nearly coherent sinusoidal or solitary zonal flows. The flows are geodesic acoustic modes (GAM), which are primarily driven by the Stringer-Winsor term. The flow amplitude together with the average anomalous transport sensitively depend on the GAM frequency and on the magnetic curvature acting on the flows, which could be influenced in a real tokamak, e.g., by shaping the plasma cross section. The local modulation of the turbulence by the flows and the excitation of the flows are due to wave-kinetic effects, which have been studied for the first time in a turbulence simulation.Comment: 5 pages, 5 figures, submitted to PR

A freely relaxing polymer remembers how it was straightened

The relaxation of initially straight semiflexible polymers has been discussed mainly with respect to the longest relaxation time. The biologically relevant non-equilibrium dynamics on shorter times is comparatively poorly understood, partly because "initially straight" can be realized in manifold ways. Combining Brownian dynamics simulations and systematic theory, we demonstrate how different experimental preparations give rise to specific short-time and universal long-time dynamics. We also discuss boundary effects and the onset of the stretch--coil transition.Comment: 14 pages, 5 figures, 3 table

The rate of beneficial mutations surfing on the wave of a range expansion

Many theoretical and experimental studies suggest that range expansions can have severe consequences for the gene pool of the expanding population. Due to strongly enhanced genetic drift at the advancing frontier, neutral and weakly deleterious mutations can reach large frequencies in the newly colonized regions, as if they were surfing the front of the range expansion. These findings raise the question of how frequently beneficial mutations successfully surf at shifting range margins, thereby promoting adaptation towards a range-expansion phenotype. Here, we use individual-based simulations to study the surfing statistics of recurrent beneficial mutations on wave-like range expansions in linear habitats. We show that the rate of surfing depends on two strongly antagonistic factors, the probability of surfing given the spatial location of a novel mutation and the rate of occurrence of mutations at that location. The surfing probability strongly increases towards the tip of the wave. Novel mutations are unlikely to surf unless they enjoy a spatial head start compared to the bulk of the population. The needed head start is shown to be proportional to the inverse fitness of the mutant type, and only weakly dependent on the carrying capacity. The second factor is the mutation occurrence which strongly decreases towards the tip of the wave. Thus, most successful mutations arise at an intermediate position in the front of the wave. We present an analytic theory for the tradeoff between these factors that allows to predict how frequently substitutions by beneficial mutations occur at invasion fronts. We find that small amounts of genetic drift increase the fixation rate of beneficial mutations at the advancing front, and thus could be important for adaptation during species invasions.Comment: 21 pages, 7 figures; to appear in PLoS Computational Biolog

Selective Sweeps in Growing Microbial Colonies

Evolutionary experiments with microbes are a powerful tool to study mutations and natural selection. These experiments, however, are often limited to the well-mixed environments of a test tube or a chemostat. Since spatial organization can significantly affect evolutionary dynamics, the need is growing for evolutionary experiments in spatially structured environments. The surface of a Petri dish provides such an environment, but a more detailed understanding of microbial growth on Petri dishes is necessary to interpret such experiments. We formulate a simple deterministic reaction-diffusion model, which successfully predicts the spatial patterns created by two competing species during colony expansion. We also derive the shape of these patterns analytically without relying on microscopic details of the model. In particular, we find that the relative fitness of two microbial strains can be estimated from the logarithmic spirals created by selective sweeps. The theory is tested with strains of the budding yeast Saccharomyces cerevisiae, for spatial competitions with different initial conditions and for a range of relative fitnesses. The reaction-diffusion model also connects the microscopic parameters like growth rates and diffusion constants with macroscopic spatial patterns and predicts the relationship between fitness in liquid cultures and on Petri dishes, which we confirmed experimentally. Spatial sector patterns therefore provide an alternative fitness assay to the commonly used liquid culture fitness assays.Molecular and Cellular Biolog

Population dynamics in compressible flows

Organisms often grow, migrate and compete in liquid environments, as well as on solid surfaces. However, relatively little is known about what happens when competing species are mixed and compressed by fluid turbulence. In these lectures we review our recent work on population dynamics and population genetics in compressible velocity fields of one and two dimensions. We discuss why compressible turbulence is relevant for population dynamics in the ocean and we consider cases both where the velocity field is turbulent and when it is static. Furthermore, we investigate populations in terms of a continuos density field and when the populations are treated via discrete particles. In the last case we focus on the competition and fixation of one species compared to anotherComment: 16 pages, talk delivered at the Geilo Winter School 201

Turbulence regulation and stabilization by equilibrium and Time-varying sheared turbulence flows

Turbulence flows are directly measured in a tokamak plasma by applying time-delay-estimation (TDE) analysis to localized 2-D density fluctuation measurements obtained with Beam Emission Spectroscopy on DIII-D. The equilibrium radial flow shear near the plasma edge (0.8 < r/a < 1) varies strongly with magnetic geometry. With the ion grad-B drift directed towards the X-point in a single null plasma, a large radial shear in the poloidal flow is measured, while little shear is observed in the reverse condition. This large shear appears to facilitate the L-to H-mode transition, consistent with the significantly lower LH transition power threshold in this configuration. In addition, time varying, radially localized (k . ρI < 1) flows with a semi-coherent structure peaked near 15 KHz and a very long poloidal wavelength, possibly m=0, are observed. These characteristics are very similar to theoretically predicted zonal flows that are self-generated by and in turn regulate the turbulence

Bacterial Filamentation Drives Colony Chirality

This is the final version. Available on open access from the American Society for Microbiology via the DOI in this recordChirality is ubiquitous in nature, with consequences at the cellular and tissue scales. As Escherichia coli colonies expand radially, an orthogonal component of growth creates a pinwheel-like pattern that can be revealed by fluorescent markers. To elucidate the mechanistic basis of this colony chirality, we investigated its link to left-handed, single-cell twisting during E. coli elongation. While chemical and genetic manipulation of cell width altered single-cell twisting handedness, colonies ceased to be chiral rather than switching handedness, and anaerobic growth altered colony chirality without affecting single-cell twisting. Chiral angle increased with increasing temperature even when growth rate decreased. Unifying these findings, we discovered that colony chirality was associated with the propensity for cell filamentation. Inhibition of cell division accentuated chirality under aerobic growth and generated chirality under anaerobic growth. Thus, regulation of cell division is intrinsically coupled to colony chirality, providing a mechanism for tuning macroscale spatial patterning. IMPORTANCE Chiral objects, such as amino acids, are distinguishable from their mirror image. For living systems, the fundamental mechanisms relating cellular handedness to chirality at the multicellular scale remain largely mysterious. Here, we use chemical, genetic, and environmental perturbations of Escherichia coli to investigate whether pinwheel patterns in bacterial colonies are directly linked to single-cell growth behaviors. We discover that chirality can be abolished without affecting single-cell twisting; instead, the degree of chirality was linked to the proportion of highly elongated cells at the colony edge. Inhibiting cell division boosted the degree of chirality during aerobic growth and even introduced chirality to otherwise achiral colonies during anaerobic growth. These findings reveal a fascinating connection between cell division and macroscopic colony patterning.National Institutes of Health (NIH)Allen Discovery Center at Stanford University on Systems Modeling of Infectio

Quasi-stationary regime of a branching random walk in presence of an absorbing wall

A branching random walk in presence of an absorbing wall moving at a constant velocity $v$ undergoes a phase transition as the velocity $v$ of the wall varies. Below the critical velocity $v_c$, the population has a non-zero survival probability and when the population survives its size grows exponentially. We investigate the histories of the population conditioned on having a single survivor at some final time $T$. We study the quasi-stationary regime for $v<v_c$ when $T$ is large. To do so, one can construct a modified stochastic process which is equivalent to the original process conditioned on having a single survivor at final time $T$. We then use this construction to show that the properties of the quasi-stationary regime are universal when $v\to v_c$. We also solve exactly a simple version of the problem, the exponential model, for which the study of the quasi-stationary regime can be reduced to the analysis of a single one-dimensional map.Comment: 2 figures, minor corrections, one reference adde

Resource limitation drives spatial organization in microbial groups.

Dense microbial groups such as bacterial biofilms commonly contain a diversity of cell types that define their functioning. However, we have a limited understanding of what maintains, or purges, this diversity. Theory suggests that resource levels are key to understanding diversity and the spatial arrangement of genotypes in microbial groups, but we need empirical tests. Here we use theory and experiments to study the effects of nutrient level on spatio-genetic structuring and diversity in bacterial colonies. Well-fed colonies maintain larger well-mixed areas, but they also expand more rapidly compared with poorly-fed ones. Given enough space to expand, therefore, well-fed colonies lose diversity and separate in space over a similar timescale to poorly fed ones. In sum, as long as there is some degree of nutrient limitation, we observe the emergence of structured communities. We conclude that resource-driven structuring is central to understanding both pattern and process in diverse microbial communities