Stochastic Neural Field Research Articles

Stochastic neural fields as gradient dynamical systems.

Continuous attractor neural networks are used extensively to model a variety of experimentally observed coherent brain states, ranging from cortical waves of activity to stationary activity bumps. The latter are thought to play an important role in various forms of neural information processing, including population coding in primary visual cortex (V1) and working memory in prefrontal cortex. However, one limitation of continuous attractor networks is that the location of the peak of an activity bump (or wave) can diffuse due to intrinsic network noise. This reflects marginal stability of bump solutions with respect to the action of an underlying continuous symmetry group. Previous studies have used perturbation theory to derive an approximate stochastic differential equation for the location of the peak (phase) of the bump. Although this method captures the diffusive wandering of a bump solution, it ignores fluctuations in the amplitude of the bump. In this paper, we show how amplitude fluctuations can be analyzed by reducing the underlying stochastic neural field equation to a finite-dimensional stochastic gradient dynamical system that tracks the stochastic motion of both the amplitude and phase of bump solutions. This allows us to derive exact expressions for the steady-state probability density and its moments, which are then used to investigate two major issues: (i) the input-dependent suppression of neural variability and (ii) noise-induced transitions to bump extinction. We develop the theory by considering the particular example of a ring attractor network with SO(2) symmetry, which is the most common architecture used in attractor models of working memory and population tuning in V1. However, we also extend the analysis to a higher-dimensional spherical attractor network with SO(3) symmetry which has previously been proposed as a model of orientation and spatial frequency tuning in V1. We thus establish how a combination of stochastic analysis and group theoretic methods provides a powerful tool for investigating the effects of noise in continuous attractor networks.

A gradient flow formulation for the stochastic Amari neural field model.

We study stochastic Amari-type neural field equations, which are mean-field models for neural activity in the cortex. We prove that under certain assumptions on the coupling kernel, the neural field model can be viewed as a gradient flow in a nonlocal Hilbert space. This makes all gradient flow methods available for the analysis, which could previously not be used, as it was not known, whether a rigorous gradient flow formulation exists. We show that the equation is well-posed in the nonlocal Hilbert space in the sense that solutions starting in this space also remain in it for all times and space-time regularity results hold for the case of spatially correlated noise. Uniqueness of invariant measures, ergodic properties for the associated Feller semigroups, and several examples of kernels are also discussed.

Stochastic neural field theory of wandering bumps on a sphere

We use a combination of group theoretic and perturbation methods to analyze the stochastic wandering of bump solutions in a neural field model on the sphere S2. We first construct an explicit bump solution in the absence of external inputs and noise, by taking the synaptic weight distribution to be the sum of first-order spherical harmonics. The corresponding neural field equation is equivariant under the action of the special orthogonal group SO(3), which implies that the bump is marginally stable with respect to rotations of the sphere. We then carry out an amplitude–phase decomposition of the solution in the presence of a weakly biased external input and weak noise, and use this to derive a pair of stochastic differential equations for the wandering of the bump, expressed in terms of angular coordinates on the sphere. The stochastic dynamics is a non-trivial generalization of the corresponding phase dynamics describing the wandering of a bump on a ring network with SO(2) symmetry, since SO(3) is non-abelian and S2 is a curved manifold.

Stochastic neural field model of stimulus-dependent variability in cortical neurons.

We use stochastic neural field theory to analyze the stimulus-dependent tuning of neural variability in ring attractor networks. We apply perturbation methods to show how the neural field equations can be reduced to a pair of stochastic nonlinear phase equations describing the stochastic wandering of spontaneously formed tuning curves or bump solutions. These equations are analyzed using a modified version of the bivariate von Mises distribution, which is well-known in the theory of circular statistics. We first consider a single ring network and derive a simple mathematical expression that accounts for the experimentally observed bimodal (or M-shaped) tuning of neural variability. We then explore the effects of inter-network coupling on stimulus-dependent variability in a pair of ring networks. These could represent populations of cells in two different layers of a cortical hypercolumn linked via vertical synaptic connections, or two different cortical hypercolumns linked by horizontal patchy connections within the same layer. We find that neural variability can be suppressed or facilitated, depending on whether the inter-network coupling is excitatory or inhibitory, and on the relative strengths and biases of the external stimuli to the two networks. These results are consistent with the general observation that increasing the mean firing rate via external stimuli or modulating drives tends to reduce neural variability.

Well-posedness of the stochastic neural field equation with discontinuous firing rate

We study the existence and uniqueness of mild solutions to the deterministic and the stochastic neural field equation with Heaviside firing rate. Since standard well-posedness results do not apply in case of a discontinuous firing rate, we present a monotone Picard iteration scheme to show the existence of a maximal mild solution. Further, we illustrate that general uniqueness does not hold, and therefore investigate uniqueness under suitable additional properties of the solutions. Here a novel criterion, the so-called absolute continuity condition is introduced. Moreover, we observe regularisation by noise: with a suitable choice of spatially correlated additive noise uniqueness is restored without imposing any additional structural assumptions.

A multiscale-analysis of stochastic bistable reaction–diffusion equations

A multiscale analysis of 1D stochastic bistable reaction–diffusion equations with additive noise is carried out w.r.t. travelling waves within the variational approach to stochastic partial differential equations. It is shown with explicit error estimates on appropriate function spaces that up to lower order w.r.t. the noise amplitude, the solution can be decomposed into the orthogonal sum of a travelling wave moving with random speed and into Gaussian fluctuations. A stochastic differential equation describing the speed of the travelling wave and a linear stochastic partial differential equation describing the fluctuations are derived in terms of the coefficients. Our results extend corresponding results obtained for stochastic neural field equations to the present class of stochastic dynamics.

Finite-Size Effects on Traveling Wave Solutions to Neural Field Equations

Neural field equations are used to describe the spatio-temporal evolution of the activity in a network of synaptically coupled populations of neurons in the continuum limit. Their heuristic derivation involves two approximation steps. Under the assumption that each population in the network is large, the activity is described in terms of a population average. The discrete network is then approximated by a continuum. In this article we make the two approximation steps explicit. Extending a model by Bressloff and Newby, we describe the evolution of the activity in a discrete network of finite populations by a Markov chain. In order to determine finite-size effects—deviations from the mean-field limit due to the finite size of the populations in the network—we analyze the fluctuations of this Markov chain and set up an approximating system of diffusion processes. We show that a well-posed stochastic neural field equation with a noise term accounting for finite-size effects on traveling wave solutions is obtained as the strong continuum limit.

Ghosts of bump attractors in stochastic neural fields: Bottlenecks and extinction

We study the dynamics of stationary bumps in continuum neural field equations near a saddle-node bifurcation. The integral terms of these evolution equations have a weight kernel describing synaptic interactions between neurons at different locations of the network. Excited regions of the neural field correspond to parts of the domain whose fraction of active neurons exceeds a sharp threshold of a firing rate nonlinearity. For sufficiently low firing threshold, a stable bump coexists with an unstable bump and a homogeneous quiescent state. As the threshold is increased, the stable and unstable branch of bump solutions annihilate in a saddle-node bifurcation. Near this criticality, we derive a quadratic amplitude equation that describes the slow evolution of the even mode (bump contractions) as it depends on the distance from the bifurcation. Beyond the bifurcation, bumps eventually become extinct, and the lifetime of bumps increases for systems nearer the bifurcation. When noise is incorporated, a stochastic amplitude equation for the even mode can be derived, which can be analyzed to quantify bump extinction time both below and above the saddle-node.

A General Framework for Stochastic Traveling Waves and Patterns, with Application to Neural Field Equations

In this paper we present a general framework in which to rigorously study the effect of spatio-temporal noise on traveling waves and stationary patterns. In particular, the framework can incorporate versions of the stochastic neural field equation that may exhibit traveling fronts, pulses, or stationary patterns. To do this, we first formulate a local SDE that describes the position of the stochastic wave up until a discontinuity time, at which point the position of the wave may jump. We then study the local stability of this stochastic front, obtaining a result that recovers a well-known deterministic result in the small-noise limit. We finish with a study of the long-time behavior of the stochastic wave.

Nonlinear Langevin Equations for Wandering Patterns in Stochastic Neural Fields

We analyze the effects of additive, spatially extended noise on spatiotemporal patterns in continuum neural fields. Our main focus is how fluctuations impact patterns when they are weakly coupled to an external stimulus or another equivalent pattern. Showing the generality of our approach, we study both propagating fronts and stationary bumps. Using a separation of time scales, we represent the effects of noise in terms of a phase-shift of a pattern from its uniformly translating position at long time scales, and fluctuations in the pattern profile around its instantaneous position at short time scales. In the case of a stimulus-locked front, we show that the phase-shift satisfies a nonlinear Langevin equation (SDE) whose deterministic part has a unique stable fixed point. Using a linear-noise approximation, we thus establish that wandering of the front about the stimulus-locked state is given by an Ornstein--Uhlenbeck (OU) process. Analogous results hold for the relative phase-shift between a pair of mutually coupled fronts, provided that the coupling is excitatory. On the other hand, if the mutual coupling is given by a Mexican hat function (difference of exponentials), the linear-noise approximation breaks down due to the coexistence of stable and unstable phase-locked states in the deterministic limit. Similarly, the stochastic motion of mutually coupled bumps can be described by a system of nonlinearly coupled SDEs, which can be linearized to yield a multivariate OU process. As in the case of fronts, large deviations can cause bumps to temporarily decouple, leading to a phase-slip in the bump positions.

Stochastic neural field equations: a rigorous footing.

We here consider a stochastic version of the classical neural field equation that is currently actively studied in the mathematical neuroscience community. Our goal is to present a well-known rigorous probabilistic framework in which to study these equations in a way that is accessible to practitioners currently working in the area, and thus to bridge some of the cultural/scientific gaps between probability theory and mathematical biology. In this way, the paper is intended to act as a reference that collects together relevant rigorous results about notions of solutions and well-posedness, which although may be straightforward to experts from SPDEs, are largely unknown in the neuroscientific community, and difficult to find in a very large body of literature. Moreover, in the course of our study we provide some new specific conditions on the parameters appearing in the equation (in particular on the neural field kernel) that guarantee the existence of a solution.

Coupling layers regularizes wave propagation in stochastic neural fields.

We explore how layered architectures influence the dynamics of stochastic neural field models. Our main focus is how the propagation of waves of neural activity in each layer is affected by interlaminar coupling. Synaptic connectivities within and between each layer are determined by integral kernels of an integrodifferential equation describing the temporal evolution of neural activity. Excitatory neural fields, with purely positive connectivities, support traveling fronts in each layer, whose speeds are increased when coupling between layers is considered. Studying the effects of noise, we find coupling reduces the variance in the position of traveling fronts, as long as the noise sources to each layer are not completely correlated. Neural fields with asymmetric connectivity support traveling pulses whose speeds are decreased by interlaminar coupling. Again, coupling reduces the variance in traveling pulse position. Asymptotic analysis is performed using a small-noise expansion, assuming interlaminar connectivity scales similarly.

Pulse Bifurcations in Stochastic Neural Fields

We study the effects of additive noise on traveling pulse solutions in spatially extended neural fields with linear adaptation. Neural fields are evolution equations with an integral term characterizing synaptic interactions between neurons at different spatial locations of the network. We introduce an auxiliary variable to model the effects of local negative feedback and consider random fluctuations by modeling the system as a set of spatially extended Langevin equations whose noise term is a $Q$-Wiener process. Due to the translation invariance of the network, neural fields can support a continuum of spatially localized bump solutions that can be destabilized by increasing the strength of the adaptation, giving rise to traveling pulse solutions. Near this criticality, we derive a stochastic amplitude equation describing the dynamics of these bifurcating pulses when the noise and the deterministic instability are of comparable magnitude. Away from this bifurcation, we investigate the effects of additive ...

Front Propagation in Stochastic Neural Fields: A Rigorous Mathematical Framework

We develop a complete and rigorous mathematical framework for the analysis of stochastic neural field equations under the influence of spatially extended additive noise. By comparing a solution to a fixed deterministic front profile it is possible to realise the difference as strong solution to an $L^2(\mathbb{R})$-valued SDE. A multiscale analysis of this process then allows us to obtain rigorous stability results. Here a new representation formula for stochastic convolutions in the semigroup approach to linear function-valued SDE with adapted random drift is applied. Additionally, we introduce a dynamic phase-adaption process of gradient type.

Large deviations for nonlocal stochastic neural fields.

We study the effect of additive noise on integro-differential neural field equations. In particular, we analyze an Amari-type model driven by a Q-Wiener process, and focus on noise-induced transitions and escape. We argue that proving a sharp Kramers’ law for neural fields poses substantial difficulties, but that one may transfer techniques from stochastic partial differential equations to establish a large deviation principle (LDP). Then we demonstrate that an efficient finite-dimensional approximation of the stochastic neural field equation can be achieved using a Galerkin method and that the resulting finite-dimensional rate function for the LDP can have a multiscale structure in certain cases. These results form the starting point for an efficient practical computation of the LDP. Our approach also provides the technical basis for further rigorous study of noise-induced transitions in neural fields based on Galerkin approximations.Mathematics Subject Classification (2000): 60F10, 60H15, 65M60, 92C20.

The effects of noise on binocular rivalry waves: a stochastic neural field model

We analyze the effects of extrinsic noise on traveling waves of visual perception in a competitive neural field model of binocular rivalry. The model consists of two one-dimensional excitatory neural fields, whose activity variables represent the responses to left-eye and right-eye stimuli, respectively. The two networks mutually inhibit each other, and slow adaptation is incorporated into the model by taking the network connections to exhibit synaptic depression. We first show how, in the absence of any noise, the system supports a propagating composite wave consisting of an invading activity front in one network co-moving with a retreating front in the other network. Using a separation of time scales and perturbation methods previously developed for stochastic reaction–diffusion equations, we then show how extrinsic noise in the activity variables leads to a diffusive-like displacement (wandering) of the composite wave from its uniformly translating position at long time scales, and fluctuations in the wave profile around its instantaneous position at short time scales. We use our analysis to calculate the first-passage-time distribution for a stochastic rivalry wave to travel a fixed distance, which we find to be given by an inverse Gaussian. Finally, we investigate the effects of noise in the depression variables, which under an adiabatic approximation lead to quenched disorder in the neural fields during propagation of a wave.

Wandering Bumps in Stochastic Neural Fields

We study the effects of noise on stationary pulse solutions (bumps) in spatially extended neural fields. The dynamics of a neural field is described by an integrodifferential equation whose integral term characterizes synaptic interactions between neurons in different spatial locations of the network. Translationally symmetric neural fields support a continuum of stationary bump solutions, which may be centered at any spatial location. Random fluctuations are introduced by modeling the system as a spatially extended Langevin equation whose noise term we take to be additive. For nonzero noise, bumps are shown to wander about the domain in a purely diffusive way. We can approximate the associated diffusion coefficient using a small noise expansion. Upon breaking the (continuous) translation symmetry of the system using spatially heterogeneous inputs or synapses, bumps in the stochastic neural field can become temporarily pinned to a finite number of locations in the network. As a result, the effective diffusion of the bump is reduced, in comparison to the homogeneous case. As the modulation frequency of this heterogeneity increases, the effective diffusion of bumps in the network approaches that of the network with spatially homogeneous weights.

Interareal coupling reduces encoding variability in multi-area models of spatial working memory.

Persistent activity observed during delayed-response tasks for spatial working memory (Funahashi et al., 1989) has commonly been modeled by recurrent networks whose dynamics is described as a bump attractor (Compte et al., 2000). We examine the effects of interareal architecture on the dynamics of bump attractors in stochastic neural fields. Lateral inhibitory synaptic structure in each area sustains stationary bumps in the absence of noise. Introducing noise causes bumps in individual areas to wander as a Brownian walk. However, coupling multiple areas together can help reduce the variability of the bump's position in each area. To examine this quantitatively, we approximate the position of the bump in each area using a small noise expansion that also assumes weak amplitude interareal projections. Our asymptotic results show the motion of the bumps in each area can be approximated as a multivariate Ornstein–Uhlenbeck process. This shows reciprocal coupling between areas can always reduce variability, if sufficiently strong, even if one area contains much more noise than the other. However, when noise is correlated between areas, the variability-reducing effect of interareal coupling is diminished. Our results suggest that distributing spatial working memory representations across multiple, reciprocally-coupled brain areas can lead to noise cancelation that ultimately improves encoding.

Laws of Large Numbers and Langevin Approximations for Stochastic Neural Field Equations

In this study, we consider limit theorems for microscopic stochastic models of neural fields. We show that the Wilson–Cowan equation can be obtained as the limit in uniform convergence on compacts in probability for a sequence of microscopic models when the number of neuron populations distributed in space and the number of neurons per population tend to infinity. This result also allows to obtain limits for qualitatively different stochastic convergence concepts, e.g., convergence in the mean. Further, we present a central limit theorem for the martingale part of the microscopic models which, suitably re-scaled, converges to a centred Gaussian process with independent increments. These two results provide the basis for presenting the neural field Langevin equation, a stochastic differential equation taking values in a Hilbert space, which is the infinite-dimensional analogue of the chemical Langevin equation in the present setting. On a technical level, we apply recently developed law of large numbers and central limit theorems for piecewise deterministic processes taking values in Hilbert spaces to a master equation formulation of stochastic neuronal network models. These theorems are valid for processes taking values in Hilbert spaces, and by this are able to incorporate spatial structures of the underlying model.Mathematics Subject Classification (2000): 60F05, 60J25, 60J75, 92C20.

Mean-field equations for stochastic firing-rate neural fields with delays: Derivation and noise-induced transitions

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.