Entropy production and coarse-graining in Markov processes

We study the large time fluctuations of entropy production in Markov processes. In particular, we consider the effect of a coarse-graining procedure which decimates {\em fast states} with respect to a given time threshold. Our results provide strong evidence that entropy production is not directly affected by this decimation, provided that it does not entirely remove loops carrying a net probability current. After the study of some examples of random walks on simple graphs, we apply our analysis to a network model for the kinesin cycle, which is an important biomolecular motor. A tentative general theory of these facts, based on Schnakenberg's network theory, is proposed.Comment: 18 pages, 13 figures, submitted for publicatio

Entropy production and coarse-graining in Markov processes

We study the large time fluctuations of entropy production in Markov processes. In particular, we consider the effect of a coarse-graining procedure which decimates {\em fast states} with respect to a given time threshold. Our results provide strong evidence that entropy production is not directly affected by this decimation, provided that it does not entirely remove loops carrying a net probability current. After the study of some examples of random walks on simple graphs, we apply our analysis to a network model for the kinesin cycle, which is an important biomolecular motor. A tentative general theory of these facts, based on Schnakenberg's network theory, is proposed.Comment: 18 pages, 13 figures, submitted for publicatio

Species lifetime distribution for simple models of ecologies

Interpretation of empirical results based on a taxa's lifetime distribution shows apparently conflicting results. Species' lifetime is reported to be exponentially distributed, whereas higher order taxa, such as families or genera, follow a broader distribution, compatible with power law decay. We show that both these evidences are consistent with a simple evolutionary model that does not require specific assumptions on species interaction. The model provides a zero-order description of the dynamics of ecological communities and its species lifetime distribution can be computed exactly. Different behaviors are found: an initial $t^{-3/2}$ power law, emerging from a random walk type of dynamics, which crosses over to a steeper $t^{-2}$ branching process-like regime and finally is cutoff by an exponential decay which becomes weaker and weaker as the total population increases. Sampling effects can also be taken into account and shown to be relevant: if species in the fossil record were sampled according to the Fisher log-series distribution, lifetime should be distributed according to a $t^{-1}$ power law. Such variability of behaviors in a simple model, combined with the scarcity of data available, cast serious doubts on the possibility to validate theories of evolution on the basis of species lifetime data.Comment: 19 pages, 2 figure

Coarse-Grained Probabilistic Automata Mimicking Chaotic Systems

Discretization of phase space usually nullifies chaos in dynamical systems. We show that if randomness is associated with discretization dynamical chaos may survive and be indistinguishable from that of the original chaotic system, when an entropic, coarse-grained analysis is performed. Relevance of this phenomenon to the problem of quantum chaos is discussed.Comment: 4 pages, 4 figure

Oscillations and temporal signalling in cells

The development of new techniques to quantitatively measure gene expression in cells has shed light on a number of systems that display oscillations in protein concentration. Here we review the different mechanisms which can produce oscillations in gene expression or protein concentration, using a framework of simple mathematical models. We focus on three eukaryotic genetic regulatory networks which show "ultradian" oscillations, with time period of the order of hours, and involve, respectively, proteins important for development (Hes1), apoptosis (p53) and immune response (NFkB). We argue that underlying all three is a common design consisting of a negative feedback loop with time delay which is responsible for the oscillatory behaviour

Anomalies, absence of local equilibrium and universality in 1-d particles systems

One dimensional systems are under intense investigation, both from theoretical and experimental points of view, since they have rather peculiar characteristics which are of both conceptual and technological interest. We analyze the dependence of the behaviour of one dimensional, time reversal invariant, nonequilibrium systems on the parameters defining their microscopic dynamics. In particular, we consider chains of identical oscillators interacting via hard core elastic collisions and harmonic potentials, driven by boundary Nos\'e-Hoover thermostats. Their behaviour mirrors qualitatively that of stochastically driven systems, showing that anomalous properties are typical of physics in one dimension. Chaos, by itslef, does not lead to standard behaviour, since it does not guarantee local thermodynamic equilibrium. A linear relation is found between density fluctuations and temperature profiles. This link and the temporal asymmetry of fluctuations of the main observables are robust against modifications of thermostat parameters and against perturbations of the dynamics.Comment: 26 pages, 16 figures, revised text, two appendices adde

Continuous coexistence or discrete species? A new review of an old question

Question: Is the coexistence of a continuum of species or ecological types possible in real-world communities? Or should one expect distinctly different species? Mathematical methods: We study whether the coexistence of species in a continuum of ecological types is (a) dynamically stable (against changes in population densities) and (b) structurally robust (against changes in population dynamics). Since most of the reviewed investigations are based on Lotka-Volterra models, we carefully explain which of the presented conclusions are model-independent. mathematical conclusions: Seemingly plausible models with dynamically stable continuous- coexistence solutions do exist. However, these models either depend on biologically unrealistic mathematical assumptions (e.g. non-differentiable ingredient functions) or are structurally unstable (i.e. destroyable by arbitrarily small modifications to those ingredient functions). The dynamical stability of a continuous-coexistence solution, if it exists, requires positive definiteness of the model's competition kernel. Biological conclusions: While the classical expectation of fixed limits to similarity is mathematically naive, the fundamental discreteness of species is a natural consequence of the basic structure of ecological interactio

Oscillation patterns in negative feedback loops

Organisms are equipped with regulatory systems that display a variety of dynamical behaviours ranging from simple stable steady states, to switching and multistability, to oscillations. Earlier work has shown that oscillations in protein concentrations or gene expression levels are related to the presence of at least one negative feedback loop in the regulatory network. Here we study the dynamics of a very general class of negative feedback loops. Our main result is that in these systems the sequence of maxima and minima of the concentrations is uniquely determined by the topology of the loop and the activating/repressing nature of the interaction between pairs of variables. This allows us to devise an algorithm to reconstruct the topology of oscillating negative feedback loops from their time series; this method applies even when some variables are missing from the data set, or if the time series shows transients, like damped oscillations. We illustrate the relevance and the limits of validity of our method with three examples: p53-Mdm2 oscillations, circadian gene expression in cyanobacteria, and cyclic binding of cofactors at the estrogen-sensitive pS2 promoter.Comment: 10 pages, 8 figure

Growth, competition and cooperation in spatial population genetics

We study an individual based model describing competition in space between two different alleles. Although the model is similar in spirit to classic models of spatial population genetics such as the stepping stone model, here however space is continuous and the total density of competing individuals fluctuates due to demographic stochasticity. By means of analytics and numerical simulations, we study the behavior of fixation probabilities, fixation times, and heterozygosity, in a neutral setting and in cases where the two species can compete or cooperate. By concluding with examples in which individuals are transported by fluid flows, we argue that this model is a natural choice to describe competition in marine environments.Comment: 29 pages, 14 figures; revised version including a section with results in the presence of fluid flow

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