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Jonathan D. Touboul

Publications and source records attributed to Jonathan D. Touboul.

13 recordsLinked to original sources

Complexity and dynamics of partially symmetric random neural networks

Neural circuits exhibit structured connectivity, including an overrepresentation of reciprocal connections between neuron pairs. Despite important advances, a full understanding of how such partial symmetry in connectivity shapes neural dynamics remains elusive. Here we ask how correlations between reciprocal connections in a random, recurrent neural network affect phase-space complexity, defined as the exponential proliferation rate (with network size) of the number of fixed points that accompanies the transition to chaotic dynamics. We find a striking pattern: partial anti-symmetry strongly amplifies complexity, while partial symmetry suppresses it. These opposing trends closely track changes in other measures of dynamical behavior, such as dimensionality, Lyapunov exponents, and transient path length, supporting the view that fixed-point structure is a key determinant of network dynamics. Thus, positive reciprocal correlations favor low-dimensional, slowly varying activity, whereas negative correlations promote high-dimensional, rapidly fluctuating chaotic activity. These results yield testable predictions about the link between connection reciprocity, neural dynamics and function.

physics.bio-ph

Pattern Formation in Mesic Savannas

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.

nlin.PS

Reliability and robustness of oscillations in some slow-fast chaotic systems

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.

nlin.CD

Spatial Dynamics with Heterogeneity

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.

math.DS

Probabilistic Foundations of Spatial Mean-field Models in Ecology and Applications

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.

math.DS

Is there sufficient evidence for criticality in cortical systems?

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.

q-bio.NC

Type III Responses to Transient Inputs in Hybrid Nonlinear Neuron Models

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.

q-bio.NC

Noise-induced synchronization and anti-resonance in excitable systems; Implications for information processing in Parkinson's Disease and Deep Brain Stimulation

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.

nlin.AO

Clamping and Synchronization in the strongly coupled FitzHugh-Nagumo model

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.

math.AP

Wild oscillations in a nonlinear neuron model with resets: (I) Bursting, spike adding and chaos

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.

math.DS

On the dynamics of random neuronal networks

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.

math.PR

Parsimony, exhaustivity and balanced detection in neocortex

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.

q-bio.NC

Canard explosion in delayed equations with multiple timescales

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.

math.DS