The mutational meltdown in asexual populations

Loss of fitness due to the accumulation of deleterious mutations appears to be inevitable in small, obligately asexual populations, as these are incapable of reconstituting highly fit genotypes by recombination or back mutation. The cumulative buildup of such mutations is expected to lead to an eventual reduction in population size, and this facilitates the chance accumulation of future mutations. This synergistic interaction between population size reduction and mutation accumulation leads to an extinction process known as the mutational meltdown, and provides a powerful explanation for the rarity of obligate asexuality. We give an overview of the theory of the mutational meltdown, showing how the process depends on the demographic properties of a population, the properties of mutations, and the relationship between fitness and number of mutations incurred

Near-periodic substitution and the genetic variance induced by environmental change

We investigate a model that describes the evolution of a diploid sexual population in a changing environment. Individuals have discrete generations and are subject to selection on the phenotypic value of a quantitative trait, which is controlled by a finite number of bialleic loci. Environmental change is taken to lead to a uniformly changing optimal phenotypic value. The population continually adapts to the changing environment, by allelic substitution, at the loci controlling the trait. We investigate the detailed interrelation between the process of allelic substitution and the adaptation and variation of the population, via infinite population calculations and finite population simulations. We find a simple relation between the substitution rate and the rate of change of the optimal phenotypic value

Temporal and dimensional effects in evolutionary graph theory

The spread in time of a mutation through a population is studied analytically and computationally in fully-connected networks and on spatial lattices. The time, t_*, for a favourable mutation to dominate scales with population size N as N^{(D+1)/D} in D-dimensional hypercubic lattices and as N ln N in fully-connected graphs. It is shown that the surface of the interface between mutants and non-mutants is crucial in predicting the dynamics of the system. Network topology has a significant effect on the equilibrium fitness of a simple population model incorporating multiple mutations and sexual reproduction. Includes supplementary information.Comment: 6 pages, 4 figures Replaced after final round of peer revie

Schwinger Boson Formulation and Solution of the Crow-Kimura and Eigen Models of Quasispecies Theory

We express the Crow-Kimura and Eigen models of quasispecies theory in a functional integral representation. We formulate the spin coherent state functional integrals using the Schwinger Boson method. In this formulation, we are able to deduce the long-time behavior of these models for arbitrary replication and degradation functions. We discuss the phase transitions that occur in these models as a function of mutation rate. We derive for these models the leading order corrections to the infinite genome length limit.Comment: 37 pages; 4 figures; to appear in J. Stat. Phy

``GLUELUMP'' SPECTRUM AND ADJOINT SOURCE POTENTIAL IN LATTICE QCD3_3

We calculate the potential between ``quarks'' which are in the adjoint representation of SU(2) color in the three-dimensional lattice theory. We work in the scaling region of the theory and at large quark separations $R$. We also calculate the masses $M_{Qg}$ of color-singlet bound states formed by coupling an adjoint quark to adjoint glue (``gluelumps''). Good scaling behavior is found for the masses of both magnetic (angular momentum $J=0$) and electric ($J=1$) gluelumps, and the magnetic gluelump is found to be the lowest-lying state. It is naively expected that the potential for adjoint quarks should saturate above a separation $R_{\rm scr}$ where it becomes energetically favorable to produce a pair of gluelumps. We obtain a good estimate of the naive screening distance $R_{\rm scr}$. However we find little evidence of saturation in the potential out to separations $R$ of about twice $R_{\rm scr}$.Comment: 8 pages plus 8 figures in 2 postscript files (uuencoded

Discharges of past flood events based on historical river profiles

International audienceThis paper presents a case study to estimate peak discharges of extreme flood events of Neckar River in south-western Germany during the 19th century. It was carried out within the BMBF research project RIMAX (Risk Management of Extreme Flood Events). The discharge estimations were made for the flood events of 1824 and 1882 based on historical cross profiles. The 1-D model Hydrologic Engineering Centers River Analysis System (HEC-RAS) was applied with different roughness coefficients. The results are compared (i) with contemporary historical calculations and (ii) in the case of a flood event in 1824 with the discharge simulation by the water balance model LARSIM (Large Area Runoff Simulation Model). These calculations are matched by the HEC-RAS simulation based on the standard roughness coefficients

Dynamics of Competitive Evolution on a Smooth Landscape

We study competitive DNA sequence evolution directed by {\it in vitro} protein binding. The steady-state dynamics of this process is well described by a shape-preserving pulse which decelerates and eventually reaches equilibrium. We explain this dynamical behavior within a continuum mean-field framework. Analytical results obtained on the motion of the pulse agree with simulations. Furthermore, finite population correction to the mean-field results are found to be insignificant.Comment: 4 pages, 2 figures, revised, to appear in Phys. Rev. Let

The statistical mechanics of a polygenic characterunder stabilizing selection, mutation and drift

By exploiting an analogy between population genetics and statistical mechanics, we study the evolution of a polygenic trait under stabilizing selection, mutation, and genetic drift. This requires us to track only four macroscopic variables, instead of the distribution of all the allele frequencies that influence the trait. These macroscopic variables are the expectations of: the trait mean and its square, the genetic variance, and of a measure of heterozygosity, and are derived from a generating function that is in turn derived by maximizing an entropy measure. These four macroscopics are enough to accurately describe the dynamics of the trait mean and of its genetic variance (and in principle of any other quantity). Unlike previous approaches that were based on an infinite series of moments or cumulants, which had to be truncated arbitrarily, our calculations provide a well-defined approximation procedure. We apply the framework to abrupt and gradual changes in the optimum, as well as to changes in the strength of stabilizing selection. Our approximations are surprisingly accurate, even for systems with as few as 5 loci. We find that when the effects of drift are included, the expected genetic variance is hardly altered by directional selection, even though it fluctuates in any particular instance. We also find hysteresis, showing that even after averaging over the microscopic variables, the macroscopic trajectories retain a memory of the underlying genetic states.Comment: 35 pages, 8 figure

Single-crossover dynamics: finite versus infinite populations

Populations evolving under the joint influence of recombination and resampling (traditionally known as genetic drift) are investigated. First, we summarise and adapt a deterministic approach, as valid for infinite populations, which assumes continuous time and single crossover events. The corresponding nonlinear system of differential equations permits a closed solution, both in terms of the type frequencies and via linkage disequilibria of all orders. To include stochastic effects, we then consider the corresponding finite-population model, the Moran model with single crossovers, and examine it both analytically and by means of simulations. Particular emphasis is on the connection with the deterministic solution. If there is only recombination and every pair of recombined offspring replaces their pair of parents (i.e., there is no resampling), then the {\em expected} type frequencies in the finite population, of arbitrary size, equal the type frequencies in the infinite population. If resampling is included, the stochastic process converges, in the infinite-population limit, to the deterministic dynamics, which turns out to be a good approximation already for populations of moderate size.Comment: 21 pages, 4 figure

20 questions on Adaptive Dynamics

Abstract Adaptive Dynamics is an approach to studying evolutionary change when fitness is density or frequency dependent. Modern papers identifying themselves as using this approach first appeared in the 1990s, and have greatly increased up to the present. However, because of the rather technical nature of many of the papers, the approach is not widely known or understood by evolutionary biologists. In this review we aim to remedy this situation by outlining the methodology and then examining its strengths and weaknesses. We carry this out by posing and answering 20 key questions on Adaptive Dynamics. We conclude that Adaptive Dynamics provides a set of useful approximations for studying various evolutionary questions. However, as with any approximate method, conclusions based on Adaptive Dynamics are valid only under some restrictions that we discuss