arXiv subjects

Michael Assaf

Publications and source records attributed to Michael Assaf.

At least 55 records · Page 3Linked to original sources

arXiv2018

Population Extinction under Bursty Reproduction in a Time Modulated Environment

In recent years non-demographic variability has been shown to greatly affect dynamics of stochastic populations. For example, non-demographic noise in the form of a bursty reproduction process with an a-priori unknown burst size, or environmental variability in the form of time-varying reaction rates, have been separately found to dramatically impact the extinction risk of isolated populations. In this work we investigate the extinction risk of an isolated population under the combined influence of these two types of non-demographic variation. Using the so-called momentum-space WKB approach we arrive at a set of time-dependent Hamilton equations. In order to account for the explicit time dependence, we find the instanton of the time-perturbed Hamiltonian numerically, where analytical expressions are presented in particular limits using various perturbation techniques. We focus on two classes of time-varying environments: periodically-varying rates corresponding to seasonal effects, and a sudden decrease in the birth rate corresponding to a catastrophe. All our theoretical results are tested against numerical Monte Carlo simulations with time-dependent rates and also against a numerical solution of the corresponding time-dependent Hamilton equations.

q-bio.PE↗

arXiv2018

Reducing the extinction risk of stochastic populations via non-demographic noise

We consider non-demographic noise in the form of uncertainty in the reaction step size, and reveal a dramatic effect this noise may have on the stability of self-regulating populations. Employing the reaction scheme mA->kA, but allowing, e.g., the product number k to be a-priori unknown and sampled from a given distribution, we show that such non-demographic noise can greatly reduce the population's extinction risk compared to the fixed k case. Our analysis is tested against numerical simulations, and by using empirical data of different species, we argue that certain distributions may be more evolutionary beneficial than others.

q-bio.PE↗

arXiv2017

Stochastic foundations in nonlinear density-regulation growth

In this work we construct individual-based models that give rise to the generalized logistic model at the mean-field deterministic level and that allow us to interpret the parameters of these models in terms of individual interactions. We also study the effect of internal fluctuations on the long-time dynamics for the different models that have been widely used in the literature, such as the theta-logistic and Savageau models. In particular, we determine the conditions for population extinction and calculate the mean time to extinction. If the population does not become extinct, we obtain analytical expressions for the population abundance distribution. Our theoretical results are based on WKB theory and the probability generating function formalism and are verified by numerical simulations.

q-bio.PE↗

arXiv2017

WKB theory of large deviations in stochastic populations

Stochasticity can play an important role in the dynamics of biologically relevant populations. These span a broad range of scales: from intra-cellular populations of molecules to population of cells and then to groups of plants, animals and people. Large deviations in stochastic population dynamics - such as those determining population extinction, fixation or switching between different states - are presently in a focus of attention of statistical physicists. We review recent progress in applying different variants of dissipative WKB approximation (after Wentzel, Kramers and Brillouin) to this class of problems. The WKB approximation allows one to evaluate the mean time and/or probability of population extinction, fixation and switches resulting from either intrinsic (demographic) noise, or a combination of the demographic noise and environmental variations, deterministic or random. We mostly cover well-mixed populations, single and multiple, but also briefly consider populations on heterogeneous networks and spatial populations. The spatial setting also allows one to study large fluctuations of the speed of biological invasions. Finally, we briefly discuss possible directions of future work.

cond-mat.stat-mech↗

arXiv2017

Large Fluctuations in Anti-Coordination Games on Scale-Free Graphs

We study the influence of the complex topology of scale-free graphs on the dynamics of anti-coordination games (e.g. snowdrift games). These reference models are characterized by the coexistence (evolutionary stable mixed strategy) of two competing species, say "cooperators" and "defectors", and, in finite systems, by metastability and large-fluctuation-driven fixation. In this work, we use extensive computer simulations and an effective diffusion approximation (in the weak selection limit) to determine under which circumstances, depending on the individual-based update rules, the topology drastically affects the long-time behavior of anti-coordination games. In particular, we compute the variance of the number of cooperators in the metastable state and the mean fixation time when the dynamics is implemented according to the voter model (death-first/birth-second process) and the link dynamics (birth/death or death/birth at random). For the voter update rule, we show that the scale-free topology effectively renormalizes the population size and as a result the statistics of observables depend on the network's degree distribution. In contrast, such a renormalization does not occur with the link dynamics update rule and we recover the same behavior as on complete graphs.

physics.soc-ph↗

arXiv2016

Colonization of a territory by a stochastic population under a strong Allee effect and a low immigration pressure

We study the dynamics of colonization of a territory by a stochastic population at low immigration pressure. We assume a sufficiently strong Allee effect that introduces, in deterministic theory, a large critical population size for colonization. At low immigration rates, the average pre-colonization population size is small thus invalidating the WKB approximation to the master equation. We circumvent this difficulty by deriving an exact zero-flux solution of the master equation and matching it with an approximate non-zero-flux solution of the pertinent Fokker-Planck equation in a small region around the critical population size. This procedure provides an accurate evaluation of the quasi-stationary probability distribution of population sizes in the pre-colonization state, and of the mean time to colonization, for a wide range of immigration rates. At sufficiently high immigration rates our results agree with WKB results obtained previously. At low immigration rates the results can be very different.

q-bio.PE↗

arXiv2016

Rare events in stochastic populations under bursty reproduction

Recently, a first step was made by the authors towards a systematic investigation of the effect of reaction-step-size noise - uncertainty in the step size of the reaction - on the dynamics of stochastic populations. This was done by investigating the effect of bursty influx on the switching dynamics of stochastic populations. Here we extend this formalism to account for bursty reproduction processes, and improve the accuracy of the formalism to include subleading-order corrections. Bursty reproduction appears in various contexts, where notable examples include bursty viral production from infected cells, and reproduction of mammals involving varying number of offspring. The main question we quantitatively address is how bursty reproduction affects the overall fate of the population. We consider two complementary scenarios: population extinction and population survival; in the former a population gets extinct after maintaining a long-lived metastable state, whereas in the latter a population proliferates despite undergoing a deterministic drift towards extinction. In both models reproduction occurs in bursts, sampled from an arbitrary distribution. In the extinction problem, we show that bursty reproduction broadens the quasi-stationary distribution of population sizes in the metastable state, which results in an exponential decrease of the mean time to extinction. In the survival problem, bursty reproduction yields an exponential increase in survival probability of the population. Close to the bifurcation limit our analytical results simplify considerably and are shown to depend solely on the mean and variance of the burst-size distribution. Our formalism is demonstrated on several realistic distributions which all compare well with numerical Monte-Carlo simulations.

cond-mat.stat-mech↗

arXiv2016

The dynamics of simple gene-network motifs subject to extrinsic fluctuations

Cellular processes do not follow deterministic rules; even in identical environments genetically identical cells can make random choices leading to different phenotypes. This randomness originates from fluctuations present in the biomolecular interaction networks. Most previous work has been focused on the intrinsic noise (IN) of these networks. Yet, especially for high-copy-number biomolecules, extrinsic or environmental noise (EN) has been experimentally shown to dominate the variation. Here, we develop an analytical formalism that allows for calculation of the effect of EN on gene-expression motifs. We introduce a method for modeling bounded EN as an auxiliary species in the master equation. The method is fully generic and is not limited to systems with small EN magnitudes. We focus our study on motifs that can be viewed as the building blocks of genetic switches: a nonregulated gene, a self-inhibiting gene, and a self-promoting gene. The role of the EN properties (magnitude, correlation time, and distribution) on the statistics of interest are systematically investigated, and the effect of fluctuations in different reaction rates is compared. Due to its analytical nature, our formalism can be used to quantify the effect of EN on the dynamics of biochemical networks and can also be used to improve the interpretation of data from single-cell gene-expression experiments.

q-bio.MN↗

arXiv2016

Effect of reaction step-size noise on the switching dynamics of stochastic populations

In genetic circuits, when the mRNA lifetime is short compared to the cell cycle, proteins are produced in geometrically-distributed bursts, which greatly affects the cellular switching dynamics between different metastable phenotypic states. Motivated by this scenario, we study a general problem of switching or escape in stochastic populations, where influx of particles occurs in groups or bursts, sampled from an arbitrary distribution. The fact that the step size of the influx reaction is a-priori unknown, and in general, may fluctuate in time with a given correlation time and statistics, introduces an additional non-demographic step-size noise into the system. Employing the probability generating function technique in conjunction with Hamiltonian formulation, we are able to map the problem in the leading order onto solving a stationary Hamilton-Jacobi equation. We show that bursty influx exponentially decreases the mean escape time compared to the "usual case" of single-step influx. In particular, close to bifurcation we find a simple analytical expression for the mean escape time, which solely depends on the mean and variance of the burst-size distribution. Our results are demonstrated on several realistic distributions and compare well with numerical Monte-Carlo simulations.

cond-mat.stat-mech↗

arXiv2015

Genetic Toggle Switch in the Absence of Cooperative Binding: Exact Results

We present an analytical treatment of a genetic switch model consisting of two mutually inhibiting genes operating without cooperative binding of the corresponding transcription factors. Previous studies have numerically shown that these systems can exhibit bimodal dynamics without possessing two stable fixed points at the deterministic level. We analytically show that bimodality is induced by the noise and find the critical repression strength that controls a transition between the bimodal and non-bimodal regimes. We also identify characteristic polynomial scaling laws of the mean switching time between bimodal states. These results, independent of the model under study, reveal essential differences between these systems and systems with cooperative binding, where there is no critical threshold for bimodality and the mean switching time scales exponentially with the system size.

q-bio.MN↗

arXiv2015

Stochastic dynamics and logistic population growth

The Verhulst model is probably the best known macroscopic rate equation in population ecology. It depends on two parameters, the intrinsic growth rate and the carrying capacity. These parameters can be estimated for different populations and are related to the reproductive fitness and the competition for limited resources, respectively. We investigate analytically and numerically the simplest possible microscopic scenarios that give rise to the logistic equation in the deterministic mean-field limit. We provide a definition of the two parameters of the Verhulst equation in terms of microscopic parameters. In addition, we derive the conditions for extinction or persistence of the population by employing either the "momentum-space" spectral theory or the "real-space" Wentzel-Kramers-Brillouin (WKB) approximation to determine the probability distribution function and the mean time to extinction of the population. Our analytical results agree well with numerical simulations.

q-bio.PE↗

arXiv2013

Cooperation dilemma in finite populations under fluctuating environments

We present a novel approach allowing the study of rare events like fixation under fluctuating environments, modeled as extrinsic noise, in evolutionary processes characterized by the dominance of one species. Our treatment consists of mapping the system onto an auxiliary model, exhibiting metastable species coexistence, that can be analyzed semiclassically. This approach enables us to study the interplay between extrinsic and demographic noise on the statistics of interest. We illustrate our theory by considering the paradigmatic prisoner's dilemma game whose evolution is described by the probability that cooperators fixate the population and replace all defectors. We analytically and numerically demonstrate that extrinsic noise may drastically enhance the cooperation fixation probability and even change its functional dependence on the population size. These results, which generalize earlier works in population genetics, indicate that extrinsic noise may help sustain and promote a much higher level of cooperation than static settings.

q-bio.PE↗

arXiv2013

Extrinsic noise driven phenotype switching in a self-regulating gene

Due to inherent noise in intracellular networks cellular decisions can be random, so genetically identical cells can display different phenotypic behavior even in identical environments. Most previous work in understanding the decision-making process has focused on the role of intrinsic noise in these systems. Yet, especially in the high copy-number regime, extrinsic noise has been shown to be much more significant. Here, using a prototypical example of a bistable self-regulating gene model, we develop a theoretical framework describing the combined effect of intrinsic and extrinsic noise on the dynamics of stochastic genetic switches. Employing our theory and Monte Carlo simulations, we show that extrinsic noise not only significantly alters the lifetimes of the phenotypic states, but can induce bistability in unexpected regions of parameter space, and may fundamentally change the escape mechanism. These results have implications for interpreting experimentally observed heterogeneity in cellular populations and for stochastic modeling of cellular decision processes.

q-bio.MN↗

arXiv2012

Effect of weak rotation on large-scale circulation cessations in turbulent convection

We investigate the effect of weak rotation on the large-scale circulation (LSC) of turbulent Rayleigh-Bénard convection, using the theory for cessations in a low-dimensional stochastic model of the flow previously studied. We determine the cessation frequency of the LSC as a function of rotation, and calculate the statistics of the amplitude and azimuthal velocity fluctuations of the LSC as a function of the rotation rate for different Rayleigh numbers. Furthermore, we show that the tails of the reorientation probability distribution function remain unchanged for rotating systems, while the distribution of the LSC amplitude and correspondingly the cessation frequency are strongly affected by rotation. Our results are in close agreement with experimental observations.

physics.flu-dyn↗

arXiv2012

Metastability and anomalous fixation in evolutionary games on scale-free networks

We study the influence of complex graphs on the metastability and fixation properties of a set of evolutionary processes. In the framework of evolutionary game theory, where the fitness and selection are frequency-dependent and vary with the population composition, we analyze the dynamics of snowdrift games (characterized by a metastable coexistence state) on scale-free networks. Using an effective diffusion theory in the weak selection limit, we demonstrate how the scale-free structure affects the system's metastable state and leads to anomalous fixation. In particular, we analytically and numerically show that the probability and mean time of fixation are characterized by stretched exponential behaviors with exponents depending on the network's degree distribution.

cond-mat.stat-mech↗

arXiv2011

Rare fluctuations and large-scale circulation cessations in turbulent convection

In turbulent Rayleigh-Benard convection, a large-scale circulation (LSC) develops in a nearly vertical plane, and is maintained by rising and falling plumes detaching from the unstable thermal boundary layers. Rare but large fluctuations in the LSC amplitude can lead to extinction of the LSC (a cessation event), followed by the re-emergence of another LSC with a different (random) azimuthal orientation. We extend previous models of the LSC dynamics to include momentum and thermal diffusion in the azimuthal plane, and calculate the tails of the probability distributions of both the amplitude and azimuthal angle. Our analytical results are in very good agreement with experimental data.

physics.flu-dyn↗

arXiv2011

Determining the stability of genetic switches: explicitly accounting for mRNA noise

Cells use genetic switches to shift between alternate gene expression states, e.g., to adapt to new environments or to follow a developmental pathway. Here, we study the dynamics of switching in a generic-feedback on/off switch. Unlike protein-only models, we explicitly account for stochastic fluctuations of mRNA, which have a dramatic impact on switch dynamics. Employing the WKB theory to treat the underlying chemical master equations, we obtain accurate results for the quasi-stationary distributions of mRNA and protein copy numbers and for the mean switching time, starting from either state. Our analytical results agree well with Monte Carlo simulations. Importantly, one can use the approach to study the effect of varying biological parameters on switch stability.

q-bio.MN↗

arXiv2011

Fixation of a Deleterious Allele under Mutation Pressure and Finite Selection Intensity

The mean fixation time of a deleterious mutant allele is studied beyond the diffusion approximation. As in Kimura's classical work [M. Kimura, Proc. Natl. Acad. Sci. U.S.A. Vol.77, 522 (1980)], that was motivated by the problem of fixation in the presence of amorphic or hypermorphic mutations, we consider a diallelic model at a single locus comprising a wild-type A and a mutant allele A' produced irreversibly from A at small uniform rate v. The relative fitnesses of the mutant homozygotes A'A', mutant heterozygotes A'A and wild-type homozygotes AA are 1-s, 1-h and 1, respectively, where it is assumed that v<< s. Here, we adopt an approach based on the direct treatment of the underlying Markov chain (birth-death process) obeyed by the allele frequency (whose dynamics is prescribed by the Moran model), which allows to accurately account for the effects of large fluctuations. After a general description of the theory, we focus on the case of a deleterious mutant allele (i.e. s>0) and discuss three situations: when the mutant is (i) completely dominant (s=h); (ii) completely recessive (h=0), and (iii) semi-dominant (h=s/2). Our theoretical predictions for the mean fixation time and the quasi-stationary distribution of the mutant population in the coexistence state, are shown to be in excellent agreement with numerical simulations. Furthermore, when s is finite, we demonstrate that our results are superior to those of the diffusion theory that is shown to be an accurate approximation only when N_e s^2 << 1, where N_e is the effective population size.

q-bio.PE↗