arXiv subjects

Nadav M. Shnerb

Publications and source records attributed to Nadav M. Shnerb.

At least 55 records · Page 3Linked to original sources

arXiv2014

A generalized neutral theory explains static and dynamic properties of biotic communities

Understanding the forces shaping ecological communities is crucially important to basic science and conservation. In recent years, considerable progress was made in explaining communities using simple and general models, with neutral theory as a prominent example. However, while successful in explaining static patterns such as species abundance distributions, the neutral theory was criticized for making unrealistic predictions of fundamental dynamic patterns. Here we incorporate environmental stochasticity into the neutral framework, and show that the resulting generalized neutral theory is capable of predicting realistic patterns of both population and community dynamics. Applying the theory to real data (the tropical forest of Barro-Colorado Island), we find that it better fits the observed distribution of short-term fluctuations, the temporal scaling of such fluctuations, and the decay of compositional similarity with time, than the original theory, while retaining its power to explain static patterns of species abundance. Importantly, although the proposed theory is neutral (all species are functionally equivalent) and stochastic, it is a niche-based theory in the sense that species differ in their demographic responses to environmental variation. Our results show that this integration of niche forces and stochasticity within a minimalistic neutral framework is highly successful in explaining fundamental static and dynamic characteristics of ecological communities.

q-bio.PE↗

arXiv2013

Stochastic Desertification

The process of desertification is usually modeled as a first order transition, where a change of an external parameter (e.g. precipitation) leads to a catastrophic bifurcation followed by an ecological regime shift. However, vegetation elements like shrubs and trees undergo a stochastic birth-death process with an absorbing state; such a process supports a second order continuous transition with no hysteresis. We present a numerical study of a minimal model that supports bistability and catastrophic shift on spatial domain with demographic noise and an absorbing state. When the external parameter varies adiabatically the transition is continuous and the front velocity renormalizes to zero at the extinction transition. Below the transition one may identify three modes of desertification: accumulation of local catastrophes, desert invasion and global collapse. A catastrophic regime shift occurs as a dynamical hysteresis, when the pace of environmental variations is too fast. We present some empirical evidence, suggesting that the mid-holocene desertification of the Sahara was, indeed, continuous.

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arXiv2013

Evolutionarily Stable Density-Dependent Dispersal

An ab-initio numerical study of the density-dependent, evolutionary stable dispersal strategy is presented. The simulations are based on a simple discretei generation island model with four processes: reproduction, dispersal, competition and local catastrophe. We do not impose any a priori constraints on the dispersal schedule, allowing the entire schedule to evolve. We find that the system converges at long times to a unique nontrivial dispersal schedule such that the dispersal probability is a monotonically increasing function of the density. We have explored the dependence of the selected dispersal strategy on the various system parameters: mean number of offspring, site carrying capacity, dispersal cost and system size. A few general scaling laws are seen to emerge from the data.

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arXiv2012

Neutral selection

Hubbell's neutral theory of biodiversity has successfully explained the observed composition of many ecological communities but it relies on strict demographic equivalence among species and provides no room for evolutionary processes like selection, adaptation and speciation. Here we show how to embed the neutral theory within the Darwinian framework. In a fitness landscape with a quadratic maximum, typical of quantitative traits, selection restricts the extant species to have traits close to optimal, so that the fitness differences between surviving species are small. For sufficiently small mutation steps, the community structure fits perfectly to the Fisher log-series species abundance distribution. The theory is relatively insensitive to moderate amounts of environmental noise, wherein the location of the fitness maximum changes by amounts of order the width of the noise-free distribution. Adding very large environmental noise to the model qualitatively changes the abundance distributions, converting the exponential fall-off of large species to a power-law decay, typical of a neutral model with environmental noise.

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arXiv2012

A scaling theory for the quasi-deterministic limit

Deterministic rate equations are widely used in the study of stochastic, interacting particles systems. This approach assumes that the inherent noise, associated with the discreteness of the elementary constituents, may be neglected when the number of particles $N$ is large. Accordingly, it fails close to the extinction transition, when the amplitude of stochastic fluctuations is comparable with the size of the population. Here we present a general scaling theory of the transition regime for spatially extended systems. Two fundamental models for out-of-equilibrium phase transitions are considered: the Susceptible-Infected-Susceptible (SIS) that belongs to the directed percolation equivalence class, and the Susceptible-Infected-Recovered (SIR) model belonging to the dynamic percolation class. Implementing the Ginzburg criteria we show that the width of the fluctuation-dominated region scales like $N^{-κ}$, where $N$ is the number of individuals per site and $κ= 2/(d_u-d)$, $d_u$ is the upper critical dimension. Other exponents that control the approach to the deterministic limit are shown to depend on $κ$. The theory is supported by the results of extensive numerical simulations for systems of various dimensionalities.

cond-mat.stat-mech↗

arXiv2011

Noise Effects in Nonlinear Biochemical Signaling

It has been generally recognized that stochasticity can play an important role in the information processing accomplished by reaction networks in biological cells. Most treatments of that stochasticity employ Gaussian noise even though it is a priori obvious that this approximation can violate physical constraints, such as the positivity of chemical concentrations. Here, we show that even when such nonphysical fluctuations are rare, an exact solution of the Gaussian model shows that the model can yield unphysical results. This is done in the context of a simple incoherent-feedforward model which exhibits perfect adaptation in the deterministic limit. We show how one can use the natural separation of time scales in this model to yield an approximate model, that is analytically solvable, including its dynamical response to an environmental change. Alternatively, one can employ a cutoff procedure to regularize the Gaussian result.

q-bio.MN↗

arXiv2010

The Birth-Death-Mutation process: a new paradigm for fat tailed distributions

Fat tailed statistics and power-laws are ubiquitous in many complex systems. Usually the appearance of of a few anomalously successful individuals (bio-species, investors, websites) is interpreted as reflecting some inherent "quality" (fitness, talent, giftedness) as in Darwin's theory of natural selection. Here we adopt the opposite, "neutral", outlook, suggesting that the main factor explaining success is merely luck. The statistics emerging from the neutral birth-death-mutation (BDM) process is shown to fit marvelously many empirical distributions. While previous neutral theories have focused on the power-law tail, our theory economically and accurately explains the entire distribution. We thus suggest the BDM distribution as a standard neutral model: effects of fitness and selection are to be identified by substantial deviations from it.

cond-mat.stat-mech↗

arXiv2010

Contact process with a defect: universal oasis, nonuniversal scaling

The extinction transition in the presence of a localized quenched defect is studied numerically. When the bulk is at criticality, the correlation length diverges and even an infinite system cannot "decouple" from the defect. The results presented here suggest that, in 1+1 dimensions, the critical exponent $δ$ that controls the asymptotic power-law decay depends on the strength of the local perturbation. On the other hand, the exponent was found to be independent of the local arrangement of the defect. In higher dimensions the defect seems to induce a transient behavior that decays algebraically in time.

cond-mat.stat-mech↗

arXiv2009

Globally coupled chaotic maps and demographic stochasticity

The affect of demographic stochasticity of a system of globally coupled chaotic maps is considered. A two-step model is studied, where the intra-patch chaotic dynamics is followed by a migration step that coupled all patches; the equilibrium number of agents on each site, $N$, controls the strength of the discreteness-induced fluctuations. For small $N$ (large fluctuations) a period-doubling cascade appears as the coupling (migration) increases. As $N$ grows an extremely slow dynamic emerges, leading to a flow along a one-dimensional family of almost period 2 solutions. This manifold become a true solutions in the deterministic limit. The degeneracy between different attractors that characterizes the clustering phase of the deterministic system is thus the $N \to \infty$ limit of the slow dynamics manifold.

nlin.CD↗

arXiv2009

Universal features of surname distribution in a subsample of a growing population

We examine the problem of family size statistics (the number of individuals carrying the same surname, or the same DNA sequence) in a given size subsample of an exponentially growing population. We approach the problem from two directions. In the first, we construct the family size distribution for the subsample from the stable distribution for the full population. This latter distribution is calculated for an arbitrary growth process in the limit of slow growth, and is seen to depend only on the average and variance of the number of children per individual, as well as the mutation rate. The distribution for the subsample is shifted left with respect to the original distribution, tending to eliminate the part of the original distribution reflecting the small families, and thus increasing the mean family size. From the subsample distribution, various bulk quantities such as the average family size and the percentage of singleton families are calculated. In the second approach, we study the past time development of these bulk quantities, deriving the statistics of the genealogical tree of the subsample. This approach reproduces that of the first when the current statistics of the subsample is considered. The surname distribution from th e 2000 U.S. Census is examined in light of these findings, and found to misrepresent the population growth rate by a factor of 1000.

q-bio.PE↗

arXiv2008

From companies to colonies: The origin of Pareto-like distributions in ecosystems

Recent studies of cluster distribution in various ecosystems revealed Pareto statistics for the size of spatial colonies. These results were supported by cellular automata simulations that yield robust criticality for endogenous pattern formation based on positive feedback. We show that this self-organized criticality is a manifestation of the law of proportionate effect, first discovered in the context of business firm size. Mapping the stochastic model to a Markov birth-death process, the transition rates are shown to scale linearly with cluster size. This mapping provides a connection between patch statistics and the dynamics of the ecosystem; the "first passage time" for different colonies emerges as a powerful tool that discriminates between endogenous and exogenous clustering mechanisms. Imminent catastrophic shifts (like desertification) manifest themselves in a drastic change of the stability properties of spatial colonies, as the chance of a cluster to disappear depends logarithmically, rather than linearly, on its size.

q-bio.PE↗

arXiv2008

Non-Hermitian delocalization and the extinction transition

Logistic growth on a static heterogenous substrate is studied both above and below the drift-induced delocalization transition. Using stochastic, agent-based simulations the delocalization of the highest eigenfunction is connected with the large $N$ limit of the stochastic theory, as the localization length of the deterministic theory controls the divergence of the spatial correlation length at the transition. Any finite colony made of discrete agents is washed away from a heterogeneity with compact support in the presence of strong wind, thus the transition belongs to the directed percolation universality class. Some of the difficulties in the analysis of the extinction transition in the presence of a localized active state are discussed.

cond-mat.stat-mech↗

arXiv2008

Extinction transition on diffusive substrate: a different universality class?

The extinction transition on a one dimensional heterogeneous substrate with diffusive correlations is studied. Diffusively correlated heterogeneity is shown to affect the location of the transition point, as the reactants adapt to the fluctuating environment. At the transition point the density decays like $t^{-0.159}$, as predicted by the theory of directed percolation. However, the scaling function describing the behavior away from the transition shows significant deviations from the DP predictions; it is suggested, thus, that the off-transition behavior of the system is governed by local adaptation to favored regions.

cond-mat.stat-mech↗

arXiv2008

Local enrichment and its nonlocal consequences for victim-exploiter metapopulations

The stabilizing effects of local enrichment are revisited. Diffusively coupled host-parasitoid and predator-prey metapopulations are shown to admit a stable fixed point, limit cycle or stable torus with a rich bifurcation structure. A linear toy model that yields many of the basic qualitative features of this system is presented. The further nonlinear complications are analyzed in the framework of the marginally stable Lotka-Volterra model, and the continuous time analog of the unstable, host-parasitoid Nicholson-Bailey model. The dependence of the results on the migration rate and level of spatial variations is examined, and the possibility of "nonlocal" effect of enrichment, where local enrichment induces stable oscillations at a distance, is studied. A simple method for basic estimation of the relative importance of this effect in experimental systems is presented and exemplified.

q-bio.PE↗

arXiv2008

Novel exponents control the quasi-deterministic limit of the extinction transition

The quasi-deterministic limit of the generic extinction transition is considered within the framework of standard epidemiological models. The susceptible-infected-susceptible (SIS) model is known to exhibit a transition from extinction to spreading, as the infectivity is increased, described by the directed percolation equivalence class. We find that the distance from the transition point, and the prefactor controlling the divergence of the (perpendicular) correlation length, both scale with the local population size, $N$, with two novel universal exponents. Different exponents characterize the large $N$ behavior of the susceptible-infected-recovered (SIR) model, which belongs to the dynamic percolation class. Extensive numerical studies in a range of systems lead to the conjecture that these characteristics are generic and may be used in order to classify the high density limit of any stochastic process on the edge of extinction.

cond-mat.stat-mech↗

arXiv2007

Desynchronization and sustainability of noisy metapopulation cycles

The apparent stability of population oscillations in ecological systems is a long-standing puzzle. A generic solution for this problem is suggested here. The stabilizing mechanism involves the combined effect of spatial migration, amplitude-dependent frequency, and noise: small differences between spatial patches, induced by the noise, are amplified by the frequency gradient. Migration among desynchronized regions then stabilizes the oscillations in the vicinity of the homogenous manifold. A simple model of diffusively coupled oscillators allows the derivation of quantitative results, like the dependence of the desynchronization upon diffusion strength and frequency differences. For an unstable system, a noise-induced transition is demonstrated, from extinction for small noise to stability if the noise exceeds some threshold. The coupled oscillator model is shown to reproduce all previously suggested stabilizing mechanisms. Accordingly, we suggest using this model as a standard tool for identification and classification of population oscillations on spatial domains.

q-bio.PE↗

arXiv2007

Solution of an infection model near threshold

We study the Susceptible-Infected-Recovered model of epidemics in the vicinity of the threshold infectivity. We derive the distribution of total outbreak size in the limit of large population size $N$. This is accomplished by mapping the problem to the first passage time of a random walker subject to a drift that increases linearly with time. We recover the scaling results of Ben-Naim and Krapivsky that the effective maximal size of the outbreak scales as $N^{2/3}$, with the average scaling as $N^{1/3}$, with an explicit form for the scaling function.

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arXiv2006

Angular velocity variations and stability of spatially explicit prey-predator systems

The linear instability of Lotka-Volterra orbits in the homogenous manifold of a two-patch system is analyzed. The origin of these orbits instability in the absence of prey migration is revealed to be the dependence of the angular velocity on the azimuthal angle; in particular, the system desynchronizes at the exit from the slow part of the trajectory. Using this insight, an analogous model of a two coupled oscillator is presented and shown to yield the same type of linear instability. This unables one to incorporate the linear instability within a recently presented general framework that allows for comparison of all known stabilization mechanisms and for simple classification of observed oscillations.

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