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We analyze a spatially extended version of a well-known model of forest-savanna dynamics, which presents as a system of nonlinear partial integro-differential equations, and study necessary conditions for pattern-forming bifurcations. Analytically, we show that homogeneous solutions dominate the dynamics of the standard forest-savanna model, regardless of the length scales of the various spatial processes considered. However, several different pattern-forming scenarios are possible upon including spatial resource limitation, such as competition for water, soil nutrients, or herbivory effects. Using numerical simulations and continuation, we study the nature of the resulting patterns as a function of system parameters and length scales, uncovering subcritical pattern-forming bifurcations and observing significant regions of multistability for realistic parameter regimes. Finally, we discuss our results in the context of extant savanna-forest modeling efforts and highlight ongoing challenges in building a unifying mathematical model for savannas across different rainfall levels., Comment: 27 pages, 7 figures

Spatial systems with heterogeneities are ubiquitous in nature, from precipitation, temperature and soil gradients controlling vegetation growth to morphogen gradients controlling gene expression in embryos. Such systems, generally described by nonlinear dynamical systems, often display complex parameter dependence and exhibit bifurcations. The dynamics of heterogeneous spatially extended systems passing through bifurcations are still relatively poorly understood, yet recent theoretical studies and experimental data highlight the resulting complex behaviors and their relevance to real-world applications. We explore the consequences of spatial heterogeneities passing through bifurcations via two examples strongly motivated by applications. These model systems illustrate that studying heterogeneity-induced behaviors in spatial systems is crucial for a better understanding of ecological transitions and functional organization in brain development., Comment: 28 pages

A variety of nonlinear models of biological systems generate complex chaotic behaviors that contrast with biological homeostasis, the observation that many biological systems prove remarkably robust in the face of changing external or internal conditions. Motivated by the subtle dynamics of cell activity in a crustacean central pattern generator (CPG), this paper proposes a refinement of the notion of chaos that reconciles homeostasis and chaos in systems with multiple timescales. We show that systems displaying relaxation cycles while going through chaotic attractors generate chaotic dynamics that are regular at macroscopic timescales and are thus consistent with physiological function. We further show that this relative regularity may break down through global bifurcations of chaotic attractors such as crises, beyond which the system may also generate erratic activity at slow timescales. We analyze these phenomena in detail in the chaotic Rulkov map, a classical neuron model known to exhibit a variety of chaotic spike patterns. This leads us to propose that the passage of slow relaxation cycles through a chaotic attractor crisis is a robust, general mechanism for the transition between such dynamics. We validate this numerically in three other models: a simple model of the crustacean CPG neural network, a discrete cubic map, and a continuous flow.

In networks of nonlinear oscillators, symmetries place hard constraints on the system that can be exploited to predict universal dynamical features and steady-states, providing a rare generic organizing principle for far-from-equilibrium systems. However, the robustness of this class of theories to symmetry-disrupting imperfections is untested. Here, we develop a model experimental reaction-diffusion network of chemical oscillators to test applications of this theory in the context of self-organizing systems relevant to biology and soft robotics. The network is a ring of 4 identical microreactors containing the oscillatory Belousov-Zhabotinsky reaction coupled to nearest neighbors via diffusion. Assuming perfect symmetry, theory predicts 4 categories of stable spatiotemporal phase-locked periodic states and 4 categories of invariant manifolds that guide and structure transitions between phase-locked states. In our experiments, we observed the predicted symmetry-derived synchronous clustered transients that occur when the dynamical trajectories coincide with invariant manifolds. However, we observe only 3 of the 4 phase-locked states that are predicted for the idealized homogeneous system. Quantitative agreement between experiment and numerical simulations is found by accounting for the small amount of experimentally determined heterogeneity. This work demonstrates that a surprising degree of the network's dynamics are constrained by symmetry in spite of the breakdown of the assumption of homogeneity and raises the question of why heterogeneity destabilizes some symmetry predicted states, but not others., Comment: *Ian and Mike contributed equally to this work. This work is under review by Physical Review X

Experimental characterization of neuronal dynamics involves recording both of spontaneous activity patterns and of responses to transient and sustained inputs. While much theoretical attention has been devoted to the spontaneous activity of neurons, less is known about the dynamic mechanisms shaping their responses to transient inputs, although these bear significant physiological relevance. Here, we study responses to transient inputs in a widely used class of neuron models (nonlinear adaptive hybrid models) well-known to reproduce a number of biologically realistic behaviors. We focus on responses to transient inputs that have been previously associated with Type III neurons, arguably the least studied category in Hodgkin's classification, which are those neurons that never exhibit continuous firing in response to sustained excitatory currents. The two phenomena that we study are post-inhibitory facilitation, in which an otherwise subthreshold excitatory input can induce a spike if it is applied with proper timing after an inhibitory pulse, and slope detection, in which a neuron spikes to a transient input only when the input's rate of change is in a specific, bounded range. We analyze the origin of these phenomena in nonlinear hybrid models and provide a geometric characterization of dynamical structures associated with PIF in the system and an analytical study of slope detection for tent inputs. While the necessary and sufficient conditions for these behaviors are easily satisfied in neurons with Type III excitability, our proofs are quite general and valid for neurons that do not exhibit Type III excitability as well. This study therefore provides a framework for the mathematical analysis of these responses to transient inputs associated with Type III neurons in other systems and for advancing our understanding of these systems' computational properties.

Many studies have found evidence that the brain operates at a critical point, a processus known as self-organized criticality. A recent paper found remarkable scalings suggestive of criticality in systems as different as neural cultures, anesthetized or awake brains. We point out here that the diversity of these states would question any claimed role of criticality in information processing. Furthermore, we show that two non-critical systems pass all the tests for criticality, a control that was not provided in the original article. We conclude that such false positives demonstrate that the presence of criticality in the brain is still not proven and that we need better methods that scaling analyses.

Deterministic models of vegetation often summarize, at a macroscopic scale, a multitude of intrinsically random events occurring at a microscopic scale. We bridge the gap between these scales by demonstrating convergence to a mean-field limit for a general class of stochastic models representing each individual ecological event in the limit of large system size. The proof relies on classical stochastic coupling techniques that we generalize to cover spatially extended interactions. The mean-field limit is a spatially extended non-Markovian process characterized by nonlocal integro-differential equations describing the evolution of the probability for a patch of land to be in a given state (the generalized Kolmogorov equations of the process, GKEs). We thus provide an accessible general framework for spatially extending many classical finite-state models from ecology and population dynamics. We demonstrate the practical effectiveness of our approach through a detailed comparison of our limiting spatial model and the finite-size version of a specific savanna-forest model, the so-called Staver-Levin model. There is remarkable dynamic consistency between the GKEs and the finite-size system, in spite of almost sure forest extinction in the finite-size system. To resolve this apparent paradox, we show that the extinction rate drops sharply when nontrivial equilibria emerge in the GKEs, and that the finite-size system's quasi-stationary distribution (stationary distribution conditional on non-extinction) closely matches the bifurcation diagram of the GKEs. Furthermore, the limit process can support periodic oscillations of the probability distribution, thus providing an elementary example of a jump process that does not converge to a stationary distribution. In spatially extended settings, environmental heterogeneity can lead to waves of invasion and front-pinning phenomena., Comment: 38 pages

We study the statistical physics of a surprising phenomenon arising in large networks of excitable elements in response to noise: while at low noise, solutions remain in the vicinity of the resting state and large-noise solutions show asynchronous activity, the network displays orderly, perfectly synchronized periodic responses at intermediate level of noise. We show that this phenomenon is fundamentally stochastic and collective in nature. Indeed, for noise and coupling within specific ranges, an asymmetry in the transition rates between a resting and an excited regime progressively builds up, leading to an increase in the fraction of excited neurons eventually triggering a chain reaction associated with a macroscopic synchronized excursion and a collective return to rest where this process starts afresh, thus yielding the observed periodic synchronized oscillations. We further uncover a novel anti-resonance phenomenon: noise-induced synchronized oscillations disappear when the system is driven by periodic stimulation with frequency within a specific range. In that anti-resonance regime, the system is optimal for measures of information capacity. This observation provides a new hypothesis accounting for the efficiency of Deep Brain Stimulation therapies in Parkinson's disease, a neurodegenerative disease characterized by an increased synchronization of brain motor circuits. We further discuss the universality of these phenomena in the class of stochastic networks of excitable elements with confining coupling, and illustrate this universality by analyzing various classical models of neuronal networks. Altogether, these results uncover some universal mechanisms supporting a regularizing impact of noise in excitable systems, reveal a novel anti-resonance phenomenon in these systems, and propose a new hypothesis for the efficiency of high-frequency stimulation in Parkinson's disease.

We investigate the dynamics of a limit of interacting FitzHugh-Nagumo neurons in the regime of large interaction coefficients. We consider the dynamics described by a mean-field model given by a nonlinear evolution partial differential equation representing the probability distribution of one given neuron in a large network. The case of weak connectivity previously studied displays a unique, exponentially stable, stationary solution. Here, we consider the case of strong connectivities, and exhibit the presence of possibly non-unique stationary behaviors or non-stationary behaviors. To this end, using Hopf-Cole transformation, we demonstrate that the solutions exponentially concentrate around a singular Dirac measure as the connectivity parameter diverges, centered at the zeros of a time-dependent continuous function. We next characterize the points at which this measure concentrates, and exhibit a particular solution corresponding to a Dirac measure concentrated on a time-dependent point satisfying an ordinary differential equation identical to the original FitzHugh-Nagumo system. This solution may thus feature multiple stable fixed points or periodic orbits, respectively corresponding to a clumping of the whole system at rest, or a synchronization of cells on a periodic solution. We illustrate these results with numerical simulations of neural networks with a relatively modest number of neurons and finite coupling strength, and show that away from the bifurcations of the limit system, the asymptotic equation recovers the main properties of more realistic networks., Comment: Added reference to the recent arxiv preprint [20]

In a series of two papers, we investigate the mechanisms by which complex oscillations are generated in a class of nonlinear dynamical systems with resets modeling the voltage and adaptation of neurons. This first paper presents mathematical analysis showing that the system can support bursts of any period as a function of model parameters. In continuous dynamical systems with resets, period-incrementing structures are complex to analyze. In the present context, we use the fact that bursting patterns correspond to periodic orbits of the adaptation map that governs the sequence of values of the adaptation variable at the resets. Using a slow-fast approach, we show that this map converges towards a piecewise linear discontinuous map whose orbits are exactly characterized. That map shows a period-incrementing structure with instantaneous transitions. We show that the period-incrementing structure persists for the full system with non-constant adaptation, but the transitions are more complex. We investigate the presence of chaos at the transitions.

In the past 20 years, the study of real eigenvalues of non-symmetric real random matrices has seen important progress. Notwithstanding, central questions still remain open, such as the characterization of their asymptotic statistics and the universality thereof. In this letter we show that for a wide class of matrices, the number $k_n$ of real eigenvalues of a matrix of size $n$ is asymptotically Gaussian with mean $\bar k_n=\mathcal{O}(\sqrt{n})$ and variance $\bar k_n(2-\sqrt{2})$. Moreover, we show that the limit distribution of real eigenvalues undergoes a transition between bimodal for $k_n=o(\sqrt{n})$ to unimodal for $k_n=\mathcal{O}(n)$, with a uniform distribution at the transition. We predict theoretically these behaviours in the Ginibre ensemble using a log-gas approach, and show numerically that they hold for a wide range of random matrices with independent entries beyond the universality class of the circular law., Comment: 10 pages, 12 figures

This work continues the analysis of large deviations for randomly connected neural networks models of the brain. The originality of the model relies on the fact that the directed impact of one particle onto another depends on the state of both particles, and (ii) have random Gaussian amplitude with mean and variance scaling as the inverse of the network size. Similarly to the spatially extended case, we show that under sufficient regularity assumptions, the empirical measure satisfies a large-deviation principle with good rate function achieving its minimum at a unique probability measure, implying in particular its convergence in both averaged and quenched cases, as well as a propagation of chaos property (in the averaged case only). The class of model we consider notably includes a stochastic version of Kuramoto model with random connections.

In a series of two papers, we investigate the large deviations and asymptotic behavior of stochastic models of brain neural networks with random interaction coefficients. In this first paper, we take into account the spatial structure of the brain and consider (i) the presence of interaction delays that depend on the distance between cells and (ii) Gaussian random interaction amplitude whose mean and variance depend on the neurons positions and scale as the inverse of the network size. We show that the empirical measure satisfies a large-deviation principle with good rate function reaching its minimum at a unique spatially extended probability measure. This result implies averaged convergence of the empirical measure and propagation of chaos. The limit is characterized through complex non-Markovian implicit equation in which the network interaction term is replaced by a non-local Gaussian process whose statistics depend on the solution over the whole neural field.

This work continues the analysis of complex dynamics in a class of bidimensional nonlinear hybrid dynamical systems with resets modeling neuronal voltage dynamics with adaptation and spike emission. We show that these models can generically display a form of mixed-mode oscillations (MMOs), which are trajectories featuring an alternation of small oscillations with spikes or bursts (multiple consecutive spikes). The mechanism by which these are generated relies fundamentally on the hybrid structure of the flow: invariant manifolds of the continuous dynamics govern small oscillations, while discrete resets govern the emission of spikes or bursts, contrasting with classical MMO mechanisms in ordinary differential equations involving more than three dimensions and generally relying on a timescale separation. The decomposition of mechanisms reveals the geometrical origin of MMOs, allowing a relatively simple classification of points on the reset manifold associated to specific numbers of small oscillations. We show that the MMO pattern can be described through the study of orbits of a discrete adaptation map, which is singular as it features discrete discontinuities with unbounded left- and right-derivatives. We study orbits of the map via rotation theory for discontinuous circle maps and elucidate in detail complex behaviors arising in the case where MMOs display at most one small oscillation between each consecutive pair of spikes.

Critical states are sometimes identified experimentally through power-law statistics or universal scaling functions. We show here that such features naturally emerge from networks in self-sustained irregular regimes away from criticality. In these regimes, statistical physics theory of large interacting systems predict a regime where the nodes have independent and identically distributed dynamics. We thus investigated the statistics of a system in which units are replaced by independent stochastic surrogates, and found the same power-law statistics, indicating that these are not sufficient to establish criticality. We rather suggest that these are universal features of large-scale networks when considered macroscopically. These results put caution on the interpretation of scaling laws found in nature., Comment: in press in Phys. Rev. E

We investigate existence and uniqueness of solutions of a McKean-Vlasov evolution PDE representing the macroscopic behaviour of interacting Fitzhugh-Nagumo neurons. This equation is hypoelliptic, nonlocal and has unbounded coefficients. We prove existence of a solution to the evolution equation and non trivial stationary solutions. Moreover, we demonstrate uniqueness of the stationary solution in the weakly nonlinear regime. Eventually, using a semigroup factorisation method, we show exponential nonlinear stability in the small connectivity regime.

We consider the ensemble of Real Ginibre matrices with a positive fraction $\alpha>0$ of real eigenvalues. We demonstrate a large deviations principle for the joint eigenvalue density of such matrices and we introduce a two phase log-gas whose stationary distribution coincides with the spectral measure of the ensemble. Using these tools we provide an asymptotic expansion for the probability $p^n_{\alpha n}$ that an $n\times n$ Ginibre matrix has $k=\alpha n$ real eigenvalues and we characterize the spectral measures of these matrices., Comment: 19 pages, 3 figures

In such different domains as neurosciences, spin glasses, social science, economics and finance, large ensemble of interacting individuals following (mainstream) or opposing (hipsters) to the majority are ubiquitous. In these systems, interactions generally occur after specific delays associated to transport, transmission or integration of information. We investigate here the impact of anti-conformism combined to delays in the emergent dynamics of large populations of mainstreams and hipsters. To this purpose, we introduce a class of simple statistical systems of interacting agents composed of (i) mainstreams and anti-conformists in the presence of (ii) delays, possibly heterogeneous, in the transmission of information. In this simple model, each agent can be in one of two states, and can change state in continuous time with a rate depending on the state of others in the past. We express the thermodynamic limit of these systems as the number of agents diverge, and investigate the solutions of the limit equation, with a particular focus on synchronized oscillations induced by delayed interactions. We show that when hipsters are too slow in detecting the trends, they will consistently make the same choice, and realizing this too late, they will switch, all together to another state where they remain alike. Similar synchronizations arise when the impact of mainstreams on hipsters choices (and reciprocally) dominate the impact of other hipsters choices, and we show that these may emerge only when the randomness in the hipsters decisions is sufficiently large. Beyond the choice of the best suit to wear this winter, this study may have important implications in understanding synchronization of nerve cells, investment strategies in finance, or emergent dynamics in social science, domains in which delays of communication and the geometry of information accessibility are prominent.

We study the mean-field limit and stationary distributions of a pulse-coupled network modeling the dynamics of a large neuronal assemblies. Our model takes into account explicitly the intrinsic randomness of firing times, contrasting with the classical integrate-and-fire model. The ergodicity properties of the Markov process associated to finite networks are investigated. We derive the limit in distribution of the sample path of the state of a neuron of the network when its size gets large. The invariant distributions of this limiting stochastic process are analyzed as well as their stability properties. We show that the system undergoes transitions as a function of the averaged connectivity parameter, and can support trivial states (where the network activity dies out, which is also the unique stationary state of finite networks in some cases) and self-sustained activity when connectivity level is sufficiently large, both being possibly stable., Comment: 37 pages, 3 figures

One fascinating aspect of the brain is its ability to process information in a fast and reliable manner. The functional architecture is thought to play a central role in this task, by encoding efficiently complex stimuli and facilitating higher level processing. In the early visual cortex of higher mammals, information is processed within functional maps whose layout is thought to underlie visual perception. The possible principles underlying the topology of the different maps, as well as the role of a specific functional architecture on information processing, is however poorly understood. We demonstrate mathematically here that two natural principles, local exhaustivity of representation and parsimony, would constrain the orientation and spatial frequency maps to display co-located singularities around which the orientation is organized as a pinwheel and spatial frequency as a dipole. This observation is perfectly in line with new optical imaging data on the cat visual cortex we analyze in a companion paper. Here we further focus on the theoretical implications of this structure. Using a computational model, we show that this architecture allows a trade-off in the local perception of orientation and spatial frequency, but this would occur for sharper selectivity than the tuning width reported in the literature. We therefore re-examined physiological data and show that indeed the spatial frequency selectivity substantially sharpens near maps singularities, bringing to the prediction that the system tends to optimize balanced detection between different attributes. These results shed new light on the principles at play in the emergence of functional architecture of cortical maps, as well as their potential role in processing information.

We analyze an inhomogeneous system of coupled reaction-diffusion equations representing the dynamics of gene expression during differentiation of nerve cells. The outcome of this developmental phase is the formation of distinct functional areas separated by sharp and smooth boundaries. It proceeds through the competition between the expression of two genes whose expression is driven by monotonic gradients of chemicals, and the products of gene expression undergo local diffusion and drive gene expression in neighboring cells. The problem therefore falls in a more general setting of species in competition within a non-homogeneous medium. We show that in the limit of arbitrarily small diffusion, there exists a unique monotonic stationary solution, which splits the neural tissue into two winner-take-all parts at a precise boundary point: on both sides of the boundary, different neuronal types are present. In order to further characterize the location of this boundary, we use a blow-up of the system and define a traveling wave problem parametrized by the position within the monotonic gradient: the precise boundary location is given by the unique point in space at which the speed of the wave vanishes.

We analyze canard explosions in delayed differential equations with a one-dimensional slow manifold. This study is applied to explore the dynamics of the van der Pol slow-fast system with delayed self-coupling. In the absence of delays, this system provides a canonical example of a canard explosion. We show that as the delay is increased a family of `classical' canard explosions ends as a Bogdanov-Takens bifurcation occurs at the folds points of the S-shaped critical manifold., Comment: arXiv admin note: substantial text overlap with arXiv:1404.5841

Motivated by the dynamics of neuronal responses, we analyze the dynamics of the Fitzhugh-Nagumo slow-fast system with delayed self-coupling. This system provides a canonical example of a canard explosion for sufficiently small delays. Beyond this regime, delays significantly enrich the dynamics, leading to mixed-mode oscillations, bursting and chaos. These behaviors emerge from a delay-induced subcritical Bogdanov-Takens instability arising at the fold points of the S-shaped critical manifold. Underlying the transition from canard-induced to delay-induced dynamics is an abrupt switch in the nature of the Hopf bifurcation.

We introduce and analyze $d$ dimensional Coulomb gases with random charge distribution and general external confining potential. We show that these gases satisfy a large deviations principle. The analysis of the minima of the rate function (which is the leading term of the energy) reveals that at equilibrium, the particle distribution is a generalized circular law (i.e. with spherical support but non-necessarily uniform distribution). In the classical electrostatic external potential, there are infinitely many minimizers of the rate function. The most likely macroscopic configuration is a disordered distribution in which particles are uniformly distributed (for $d=2$, the circular law), and charges are independent of the positions of the particles. General charge-dependent confining potentials unfold this degenerate situation: in contrast, the particle density is not uniform, and particles spontaneously organize according to their charge. In that picture the classical electrostatic potential appears as a transition at which order is lost. Sub-leading terms of the energy are derived: we show that these are related to an operator, generalizing the Coulomb renormalized energy, which incorporates the heterogeneous nature of the charges. This heterogeneous renormalized energy informs us about the microscopic arrangements of the particles, which are non-standard, strongly depending on the charges, and include progressive and irregular lattices., Comment: 26 pages, 10 figures

Neural field equations are integro-differential systems describing the macroscopic activity of spatially extended pieces of cortex. In such cortical assemblies, the propagation of information and the transmission machinery induce communication delays, due to the transport of information (propagation delays) and to the synaptic machinery (constant delays). We investigate the role of these delays on the formation of structured spatiotemporal patterns for these systems in arbitrary dimensions. We focus on localized activity, either induced by the presence of a localized stimulus (pulses) or by transitions between two levels of activity (fronts). Linear stability analysis allows to reveal the existence of Hopf bifurcation curves induced by the delays, along different modes that may be symmetric or asymmetric. We show that instabilities strongly depend on the dimension, and in particular may exhibit transversal instabilities along invariant directions. These instabilities yield pulsatile localized activity, and depending on the symmetry of the destabilized modes, either produce spatiotemporal breathing or sloshing patterns.

Complex systems, and in particular random neural networks, are often described by randomly interacting dynamical systems with no specific symmetry. In that context, characterizing the number of relevant directions necessitates fine estimates on the Ginibre ensemble. In this Letter, we compute analytically the probability distribution of the number of eigenvalues $N_R$ with modulus greater than $R$ (the index) of a large $N\times N$ random matrix in the real or complex Ginibre ensemble. We show that the fraction $N_R/N=p$ has a distribution scaling as $\exp(-\beta N^2 \psi_R(p))$ with $\beta=1$ (respectively $\beta=1/2$) for the complex (resp. real) Ginibre ensemble. For any $p\in[0,1]$, the equilibrium spectral densities as well as the rate function $\psi_R(p)$ are explicitly derived. This function displays a third order phase transition at the critical (minimum) value $p^*_R=1-R^2$, associated to a phase transition of the Coulomb gas. We deduce that, in the central regime, the fluctuations of the index $N_R$ around its typical value $p^*_R N$ scale as $N^{1/3}$., Comment: 7 pages, 2 figures

The cortex is a very large network characterized by a complex connectivity including at least two scales: a microscopic scale at which the interconnections are non-specific and very dense, while macroscopic connectivity patterns connecting different regions of the brain at larger scale are extremely sparse. This motivates to analyze the behavior of networks with multiscale coupling, in which a neuron is connected to its $v(N)$ nearest-neighbors where $v(N)=o(N)$, and in which the probability of macroscopic connection between two neurons vanishes. These are called singular multi-scale connectivity patterns. We introduce a class of such networks and derive their continuum limit. We show convergence in law and propagation of chaos in the thermodynamic limit. The limit equation obtained is an intricate non-local McKean-Vlasov equation with delays which is universal with respect to the type of micro-circuits and macro-circuits involved., Comment: arXiv admin note: text overlap with arXiv:1108.2414

Networks of the brain are composed of a very large number of neurons connected through a random graph and interacting after random delays that both depend on the anatomical distance between cells. In order to comprehend the role of these random architectures on the dynamics of such networks, we analyze the mesoscopic and macroscopic limits of networks with random correlated connectivity weights and delays. We address both averaged and quenched limits, and show propagation of chaos and convergence to a complex integral McKean-Vlasov equations with distributed delays. We then instantiate a completely solvable model illustrating the role of such random architectures in the emerging macroscopic activity. We particularly focus on the role of connectivity levels in the emergence of periodic solutions.

Characterizing the in uence of network properties on the global emerging behavior of interacting elements constitutes a central question in many areas, from physical to social sciences. In this article we study a primary model of disordered neuronal networks with excitatory-inhibitory structure and balance constraints. We show how the interplay between structure and disorder in the connectivity leads to a universal transition from trivial to synchronized stationary or periodic states. This transition cannot be explained only through the analysis of the spectral density of the connectivity matrix. We provide a low dimensional approximation that shows the role of both the structure and disorder in the dynamics., Comment: 5 pages, 3 figures

We investigate the dynamics of large stochastic networks with different timescales and nonlinear mean-field interactions. After deriving the limit equations for a general class of network models, we apply our results to the celebrated Wilson-Cowan system with two populations with or without slow adaptation, paradigmatic example of nonlinear mean-field network. This system has the property that the dynamics of the mean of the solution exactly satisfies an ODE. This reduction allows to show that in the mean-field limit and in multiple populations with multiple timescales, noise induces canard explosions and Mixed-Mode Oscillations on the mean of the solution. This sheds new light on the qualitative effects of noise and sensitivity to precise noise values in large stochastic networks. We further investigate finite-sized networks and show that systematic differences with the mean-field limits arise in bistable regimes (where random switches between different attractors occur) or in mixed-mode oscillations, were the finite-size effects induce early jumps due to the sensitivity of the attractor.

At functional scales, cortical behavior results from the complex interplay of a large number of excitable cells operating in noisy environments. Such systems resist to mathematical analysis, and computational neurosciences have largely relied on heuristic partial (and partially justified) macroscopic models, which successfully reproduced a number of relevant phenomena. The relationship between these macroscopic models and the spiking noisy dynamics of the underlying cells has since then been a great endeavor. Based on recent mean-field reductions for such spiking neurons, we present here {a principled reduction of large biologically plausible neuronal networks to firing-rate models, providing a rigorous} relationship between the macroscopic activity of populations of spiking neurons and popular macroscopic models, under a few assumptions (mainly linearity of the synapses). {The reduced model we derive consists of simple, low-dimensional ordinary differential equations with} parameters and {nonlinearities derived from} the underlying properties of the cells, and in particular the noise level. {These simple reduced models are shown to reproduce accurately the dynamics of large networks in numerical simulations}. Appropriate parameters and functions are made available {online} for different models of neurons: McKean, Fitzhugh-Nagumo and Hodgkin-Huxley models.

We analyze the macroscopic behavior of multi-populations randomly connected neural networks with interaction delays. Similar to cases occurring in spin glasses, we show that the sequences of empirical measures satisfy a large deviation principle, and converge towards a self-consistent non-Markovian process. The proof differs in that we are working in infinite-dimensional spaces (interaction delays), non-centered interactions and multiple cell types. The limit equation is qualitatively analyzed, and we identify a number of phase transitions in such systems upon changes in delays, connectivity patterns and dispersion, particularly focusing on the emergence of non-equilibrium states involving synchronized oscillations.

Random neural networks are dynamical descriptions of randomly interconnected neural units. These show a phase transition to chaos as a disorder parameter is increased. The microscopic mechanisms underlying this phase transition are unknown, and similarly to spin-glasses, shall be fundamentally related to the behavior of the system. In this Letter we investigate the explosion of complexity arising near that phase transition. We show that the mean number of equilibria undergoes a sharp transition from one equilibrium to a very large number scaling exponentially with the dimension on the system. Near criticality, we compute the exponential rate of divergence, called topological complexity. Strikingly, we show that it behaves exactly as the maximal Lyapunov exponent, a classical measure of dynamical complexity. This relationship unravels a microscopic mechanism leading to chaos which we further demonstrate on a simpler class of disordered systems, suggesting a deep and underexplored link between topological and dynamical complexity.

Realistic large-scale networks display an heterogeneous distribution of connectivity weights, that might also randomly vary in time. We show that depending on the level of heterogeneity in the connectivity coefficients, different qualitative macroscopic and microscopic regimes emerge. We evidence in particular generic transitions from stationary to perfectly periodic phase-locked regimes as the disorder parameter is increased, both in a simple model treated analytically and in a biologically relevant network made of excitable cells.

Realistic networks display heterogeneous transmission delays. We analyze here the limits of large stochastic multi-populations networks with stochastic coupling and random interconnection delays. We show that depending on the nature of the delays distributions, a quenched or averaged propagation of chaos takes place in these networks, and that the network equations converge towards a delayed McKean-Vlasov equation with distributed delays. Our approach is mostly fitted to neuroscience applications. We instantiate in particular a classical neuronal model, the Wilson and Cowan system, and show that the obtained limit equations have Gaussian solutions whose mean and standard deviation satisfy a closed set of coupled delay differential equations in which the distribution of delays and the noise levels appear as parameters. This allows to uncover precisely the effects of noise, delays and coupling on the dynamics of such heterogeneous networks, in particular their role in the emergence of synchronized oscillations. We show in several examples that not only the averaged delay, but also the dispersion, govern the dynamics of such networks., Comment: Corrected misprint (useless stopping time) in proof of Lemma 1 and clarified a regularity hypothesis (remark 1)

In this article, we address the absorption properties of a class of stochastic differ- ential equations around singular points where both the drift and diffusion functions vanish. According to the H\"older coefficient alpha of the diffusion function around the singular point, we identify different regimes. Stability of the absorbing state, large deviations for the absorption time, existence of stationary or quasi-stationary distributions are discussed. In particular, we show that quasi-stationary distributions only exist for alpha < 3/4, and for alpha in the interval (3/4, 1), no quasi-stationary distribution is found and numerical simulations tend to show that the process conditioned on not being absorbed initiates an almost sure exponential convergence towards the absorbing state (as is demonstrated to be true for alpha = 1). Applications of these results to stochastic bifurcations are discussed.

We derive the mean-field equations arising as the limit of a network of interacting spiking neurons, as the number of neurons goes to infinity. The neurons belong to a fixed number of populations and are represented either by the Hodgkin-Huxley model or by one of its simplified version, the Fitzhugh-Nagumo model. The synapses between neurons are either electrical or chemical. The network is assumed to be fully connected. The maximum conductances vary randomly. Under the condition that all neurons initial conditions are drawn independently from the same law that depends only on the population they belong to, we prove that a propagation of chaos phenomenon takes places, namely that in the mean-field limit, any finite number of neurons become independent and, within each population, have the same probability distribution. This probability distribution is solution of a set of implicit equations, either nonlinear stochastic differential equations resembling the McKean-Vlasov equations, or non-local partial differential equations resembling the McKean-Vlasov-Fokker- Planck equations. We prove the well-posedness of these equations, i.e. the existence and uniqueness of a solution. We also show the results of some preliminary numerical experiments that indicate that the mean-field equations are a good representation of the mean activity of a finite size network, even for modest sizes. These experiment also indicate that the McKean-Vlasov-Fokker- Planck equations may be a good way to understand the mean-field dynamics through, e.g., a bifurcation analysis., Comment: 55 pages, 9 figures

In this manuscript we analyze the collective behavior of mean-field limits of large-scale, spatially extended stochastic neuronal networks with delays. Rigorously, the asymptotic regime of such systems is characterized by a very intricate stochastic delayed integro-differential McKean-Vlasov equation that remain impenetrable, leaving the stochastic collective dynamics of such networks poorly understood. In order to study these macroscopic dynamics, we analyze networks of firing-rate neurons, i.e. with linear intrinsic dynamics and sigmoidal interactions. In that case, we prove that the solution of the mean-field equation is Gaussian, hence characterized by its two first moments, and that these two quantities satisfy a set of coupled delayed integro-differential equations. These equations are similar to usual neural field equations, and incorporate noise levels as a parameter, allowing analysis of noise-induced transitions. We identify through bifurcation analysis several qualitative transitions due to noise in the mean-field limit. In particular, stabilization of spatially homogeneous solutions, synchronized oscillations, bumps, chaotic dynamics, wave or bump splitting are exhibited and arise from static or dynamic Turing-Hopf bifurcations. These surprising phenomena allow further exploring the role of noise in the nervous system., Comment: Updated to the latest version published, and clarified the dependence in space of Brownian motions

We consider the problem of the limit of bio-inspired spatially extended neuronal networks including an infinite number of neuronal types (space locations), with space-dependent propagation delays modeling neural fields. The propagation of chaos property is proved in this setting under mild assumptions on the neuronal dynamics, valid for most models used in neuroscience, in a mesoscopic limit, the neural-field limit, in which we can resolve the quite fine structure of the neuron's activity in space and where averaging effects occur. The mean-field equations obtained are of a new type: they take the form of well-posed infinite-dimensional delayed integro-differential equations with a nonlocal mean-field term and a singular spatio-temporal Brownian motion. We also show how these intricate equations can be used in practice to uncover mathematically the precise mesoscopic dynamics of the neural field in a particular model where the mean-field equations exactly reduce to deterministic nonlinear delayed integro-differential equations. These results have several theoretical implications in neuroscience we review in the discussion., Comment: Updated to correct an erroneous suggestion of extension of the results in Appendix B, and to clarify some measurability questions in the proof of Theorem 2

The collective behavior of cortical neurons is strongly affected by the presence of noise at the level of individual cells. In order to study these phenomena in large-scale assemblies of neurons, we consider networks of firing-rate neurons with linear intrinsic dynamics and nonlinear coupling, belonging to a few types of cell populations and receiving noisy currents. Asymptotic equations as the number of neurons tends to infinity (mean field equations) are rigorously derived based on a probabilistic approach. These equations are implicit on the probability distribution of the solutions which generally makes their direct analysis difficult. However, in our case, the solutions are Gaussian, and their moments satisfy a closed system of nonlinear ordinary differential equations (ODEs), which are much easier to study than the original stochastic network equations, and the statistics of the empirical process uniformly converge towards the solutions of these ODEs. Based on this description, we analytically and numerically study the influence of noise on the collective behaviors, and compare these asymptotic regimes to simulations of the network. We observe that the mean field equations provide an accurate description of the solutions of the network equations for network sizes as small as a few hundreds of neurons. In particular, we observe that the level of noise in the system qualitatively modifies its collective behavior, producing for instance synchronized oscillations of the whole network, desynchronization of oscillating regimes, and stabilization or destabilization of stationary solutions. These results shed a new light on the role of noise in shaping collective dynamics of neurons, and gives us clues for understanding similar phenomena observed in biological networks.

Based on the analysis of a certain class of linear operators on a Banach space, we provide a closed form expression for the solutions of certain linear partial differential equations with non-autonomous input, time delays and stochastic terms, which takes the form of an infinite series expansion.

The study of multidimensional stochastic processes involves complex computations in intricate functional spaces. In particular, the diffusion processes, which include the practically important Gauss-Markov processes, are ordinarily defined through the theory of stochastic integration. Here, inspired by the L\'{e}vy-Cieselski construction of the Wiener process, we propose an alternative representation of multidimensional Gauss-Markov processes as expansions on well-chosen Schauder bases, with independent random coefficients of normal law with zero mean and unitary variance. We thereby offer a natural multi-resolution description of Gauss-Markov processes as limits of the finite-dimensional partial sums of the expansion, that are strongly almost-surely convergent. Moreover, such finite-dimensional random processes constitute an optimal approximation of the process, in the sense of minimizing the associated Dirichlet energy under interpolating constraints. This approach allows simpler treatment in many applied and theoretical fields and we provide a short overview of applications we are currently developing., Comment: 94 pages, 4 figures

The brain's activity is characterized by the interaction of a very large number of neurons that are strongly affected by noise. However, signals often arise at macroscopic scales integrating the effect of many neurons into a reliable pattern of activity. In order to study such large neuronal assemblies, one is often led to derive mean-field limits summarizing the effect of the interaction of a large number of neurons into an effective signal. Classical mean-field approaches consider the evolution of a deterministic variable, the mean activity, thus neglecting the stochastic nature of neural behavior. In this article, we build upon two recent approaches that include correlations and higher order moments in mean-field equations, and study how these stochastic effects influence the solutions of the mean-field equations, both in the limit of an infinite number of neurons and for large yet finite networks. We introduce a new model, the infinite model, which arises from both equations by a rescaling of the variables and, which is invertible for finite-size networks, and hence, provides equivalent equations to those previously derived models. The study of this model allows us to understand qualitative behavior of such large-scale networks. We show that, though the solutions of the deterministic mean-field equation constitute uncorrelated solutions of the new mean-field equations, the stability properties of limit cycles are modified by the presence of correlations, and additional non-trivial behaviors including periodic orbits appear when there were none in the mean field. The origin of all these behaviors is then explored in finite-size networks where interesting mesoscopic scale effects appear. This study leads us to show that the infinite-size system appears as a singular limit of the network equations, and for any finite network, the system will differ from the infinite system.

Bidimensional spiking models currently gather a lot of attention for their simplicity and their ability to reproduce various spiking patterns of cortical neurons, and are particularly used for large network simulations. These models describe the dynamics of the membrane potential by a nonlinear differential equation that blows up in finite time, coupled to a second equation for adaptation. Spikes are emitted when the membrane potential blows up or reaches a cutoff value. The precise simulation of the spike times and of the adaptation variable is critical for it governs the spike pattern produced, and is hard to compute accurately because of the exploding nature of the system at the spike times. We thoroughly study the precision of fixed time-step integration schemes for this type of models and demonstrate that these methods produce systematic errors that are unbounded, as the cutoff value is increased, in the evaluation of the two crucial quantities: the spike time and the value of the adaptation variable at this time. Precise evaluation of these quantities therefore involve very small time steps and long simulation times. In order to achieve a fixed absolute precision in a reasonable computational time, we propose here a new algorithm to simulate these systems based on a variable integration step method that either integrates the original ordinary differential equation or the equation of the orbits in the phase plane, and compare this algorithm with fixed time-step Euler scheme and other more accurate simulation algorithms.

Emotional disorders and psychological flourishing are the result of complex interactions between positive and negative affects that depend on external events and the subject's internal representations. Based on psychological data, we mathematically model the dynamical balance between positive and negative affects as a function of the response to external positive and negative events. This modeling allows the investigation of the relative impact of two leading forms of therapy on affect balance. The model uses a delay differential equation to analytically study the complete bifurcation diagram of the system. We compare the results of the model to psychological data on a single, recurrently depressed patient that was administered the two types of therapies considered (viz., coping-focused vs. affect-focused). The model leads to the prediction that stabilization at a normal state may rely on evaluating one's emotional state through an historical ongoing emotional state rather than in a narrow present window. The simple mathematical model proposed here offers a theoretically grounded quantitative framework for investigating the temporal process of change and parameters of resilience to relapse.

We review a recent approach to the mean-field limits in neural networks that takes into account the stochastic nature of input current and the uncertainty in synaptic coupling. This approach was proved to be a rigorous limit of the network equations in a general setting, and we express here the results in a more customary and simpler framework. We propose a heuristic argument to derive these equations providing a more intuitive understanding of their origin. These equations are characterized by a strong coupling between the different moments of the solutions. We analyse the equations, present an algorithm to simulate the solutions of these mean-field equations, and investigate numerically the equations. In particular, we build a bridge between these equations and Sompolinsky and collaborators approach (1988, 1990), and show how the coupling between the mean and the covariance function deviates from customary approaches.