Control of seizure-like dynamics in neuronal populations with excitability adaptation related to ketogenic diet.

We consider a heterogeneous, globally coupled population of excitatory quadratic integrate-and-fire neurons with excitability adaptation due to a metabolic feedback associated with ketogenic diet, a form of therapy for epilepsy. Bifurcation analysis of a three-dimensional mean-field system derived in the framework of next-generation neural mass models allows us to explain the scenarios and suggest control strategies for the transitions between the neurophysiologically desired asynchronous states and the synchronous, seizure-like states featuring collective oscillations. We reveal two qualitatively different scenarios for the onset of synchrony. For weaker couplings, a bistability region between the lower- and the higher-activity asynchronous states unfolds from the cusp point, and the collective oscillations emerge via a supercritical Hopf bifurcation. For stronger couplings, one finds seven co-dimension two bifurcation points, including pairs of Bogdanov-Takens and generalized Hopf points, such that both lower- and higher-activity asynchronous states undergo transitions to collective oscillations, with hysteresis and jump-like behavior observed in vicinity of subcritical Hopf bifurcations. We demonstrate three control mechanisms for switching between asynchronous and synchronous states, involving parametric perturbation of the adenosine triphosphate (ATP) production rate, external stimulation currents, or pulse-like ATP shocks, and indicate a potential therapeutic advantage of hysteretic scenarios.

Scale-free avalanches in arrays of FitzHugh-Nagumo oscillators.

The activity in the brain cortex remarkably shows a simultaneous presence of robust collective oscillations and neuronal avalanches, where intermittent bursts of pseudo-synchronous spiking are interspersed with long periods of quiescence. The mechanisms allowing for such coexistence are still a matter of an intensive debate. Here, we demonstrate that avalanche activity patterns can emerge in a rather simple model of an array of diffusively coupled neural oscillators with multiple timescale local dynamics in the vicinity of a canard transition. The avalanches coexist with the fully synchronous state where the units perform relaxation oscillations. We show that the mechanism behind the avalanches is based on an inhibitory effect of interactions, which may quench the spiking of units due to an interplay with the maximal canard. The avalanche activity bears certain heralds of criticality, including scale-invariant distributions of event sizes. Furthermore, the system shows increased sensitivity to perturbations, manifested as critical slowing down and reduced resilience.

Perspectives on adaptive dynamical systems.

Adaptivity is a dynamical feature that is omnipresent in nature, socio-economics, and technology. For example, adaptive couplings appear in various real-world systems, such as the power grid, social, and neural networks, and they form the backbone of closed-loop control strategies and machine learning algorithms. In this article, we provide an interdisciplinary perspective on adaptive systems. We reflect on the notion and terminology of adaptivity in different disciplines and discuss which role adaptivity plays for various fields. We highlight common open challenges and give perspectives on future research directions, looking to inspire interdisciplinary approaches.

Patched patterns and emergence of chaotic interfaces in arrays of nonlocally coupled excitable systems.

We disclose a new class of patterns, called patched patterns, in arrays of non-locally coupled excitable units with attractive and repulsive interactions. The self-organization process involves the formation of two types of patches, majority and minority ones, characterized by uniform average spiking frequencies. Patched patterns may be temporally periodic, quasiperiodic, or chaotic, whereby chaotic patterns may further develop interfaces comprised of units with average frequencies in between those of majority and minority patches. Using chaos and bifurcation theory, we demonstrate that chaos typically emerges via a torus breakup and identify the secondary bifurcation that gives rise to chaotic interfaces. It is shown that the maximal Lyapunov exponent of chaotic patched patterns does not decay, but rather converges to a finite value with system size. Patched patterns with a smaller wavenumber may exhibit diffusive motion of chaotic interfaces, similar to that of the incoherent part of chimeras.

Unbalanced clustering and solitary states in coupled excitable systems.

We discover the mechanisms of emergence and the link between two types of symmetry-broken states, the unbalanced periodic two-cluster states and solitary states, in coupled excitable systems with attractive and repulsive interactions. The prevalent solitary states in non-locally coupled arrays, whose self-organization is based on successive (order preserving) spiking of units, derive their dynamical features from the corresponding unbalanced cluster states in globally coupled networks. Apart from the states with successive spiking, we also find cluster and solitary states where the interplay of excitability and local multiscale dynamics gives rise to so-called leap-frog activity patterns with an alternating order of spiking between the units. We show that the noise affects the system dynamics by suppressing the multistability of cluster states and by inducing pattern homogenization, transforming solitary states into patterns of patched synchrony.

Collective Activity Bursting in a Population of Excitable Units Adaptively Coupled to a Pool of Resources.

We study the collective dynamics in a population of excitable units (neurons) adaptively interacting with a pool of resources. The resource pool is influenced by the average activity of the population, whereas the feedback from the resources to the population is comprised of components acting homogeneously or inhomogeneously on individual units of the population. Moreover, the resource pool dynamics is assumed to be slow and has an oscillatory degree of freedom. We show that the feedback loop between the population and the resources can give rise to collective activity bursting in the population. To explain the mechanisms behind this emergent phenomenon, we combine the Ott-Antonsen reduction for the collective dynamics of the population and singular perturbation theory to obtain a reduced system describing the interaction between the population mean field and the resources.

Bumps, chimera states, and Turing patterns in systems of coupled active rotators.

Self-organized coherence-incoherence patterns, called chimera states, have first been reported in systems of Kuramoto oscillators. For coupled excitable units, similar patterns where coherent units are at rest are called bump states. Here, we study bumps in an array of active rotators coupled by nonlocal attraction and global repulsion. We demonstrate how they can emerge in a supercritical scenario from completely coherent Turing patterns: a single incoherent unit appears in a homoclinic bifurcation, undergoing subsequent transitions to quasiperiodic and chaotic behavior, which eventually transforms into extensive chaos with many incoherent units. We present different types of transitions and explain the formation of coherence-incoherence patterns according to the classical paradigm of short-range activation and long-range inhibition.

Dynamics of a stochastic excitable system with slowly adapting feedback.

We study an excitable active rotator with slowly adapting nonlinear feedback and noise. Depending on the adaptation and the noise level, this system may display noise-induced spiking, noise-perturbed oscillations, or stochastic bursting. We show how the system exhibits transitions between these dynamical regimes, as well as how one can enhance or suppress the coherence resonance or effectively control the features of the stochastic bursting. The setup can be considered a paradigmatic model for a neuron with a slow recovery variable or, more generally, as an excitable system under the influence of a nonlinear control mechanism. We employ a multiple timescale approach that combines the classical adiabatic elimination with averaging of rapid oscillations and stochastic averaging of noise-induced fluctuations by a corresponding stationary Fokker-Planck equation. This allows us to perform a numerical bifurcation analysis of a reduced slow system and to determine the parameter regions associated with different types of dynamics. In particular, we demonstrate the existence of a region of bistability, where the noise-induced switching between a stationary and an oscillatory regime gives rise to stochastic bursting.

Two paradigmatic scenarios for inverse stochastic resonance.

Inverse stochastic resonance comprises a nonlinear response of an oscillatory system to noise where the frequency of noise-perturbed oscillations becomes minimal at an intermediate noise level. We demonstrate two generic scenarios for inverse stochastic resonance by considering a paradigmatic model of two adaptively coupled stochastic active rotators whose local dynamics is close to a bifurcation threshold. In the first scenario, shown for the two rotators in the excitable regime, inverse stochastic resonance emerges due to a biased switching between the oscillatory and the quasi-stationary metastable states derived from the attractors of the noiseless system. In the second scenario, illustrated for the rotators in the oscillatory regime, inverse stochastic resonance arises due to a trapping effect associated with a noise-enhanced stability of an unstable fixed point. The details of the mechanisms behind the resonant effect are explained in terms of slow-fast analysis of the corresponding noiseless systems.

Leap-frog patterns in systems of two coupled FitzHugh-Nagumo units.

We study a system of two identical FitzHugh-Nagumo units with a mutual linear coupling in the fast variables. While an attractive coupling always leads to synchronous behavior, a repulsive coupling can give rise to dynamical regimes with alternating spiking order, called leap-frogging. We analyze various types of periodic and chaotic leap-frogging regimes, using numerical path-following methods to investigate their emergence and stability, as well as to obtain the complex bifurcation scenario which organizes their appearance in parameter space. In particular, we show that the stability region of the simplest periodic leap-frog pattern has the shape of a locking cone pointing to the canard transition of the uncoupled system. We also discuss the role of the timescale separation in the coupled FitzHugh-Nagumo system and the relation of the leap-frog solutions to the theory of mixed-mode oscillations in multiple timescale systems.

Two scenarios for the onset and suppression of collective oscillations in heterogeneous populations of active rotators.

We consider the macroscopic regimes and the scenarios for the onset and the suppression of collective oscillations in a heterogeneous population of active rotators composed of excitable or oscillatory elements. We analyze the system in the continuum limit within the framework of Ott-Antonsen reduction method, determining the states with a constant mean field and their stability boundaries in terms of the characteristics of the rotators' frequency distribution. The system is established to display three macroscopic regimes, namely the homogeneous stationary state, where all the units lie at the resting state, the global oscillatory state, characterized by the partially synchronized local oscillations, and the heterogeneous stationary state, which includes a mixture of resting and asynchronously oscillating units. The transitions between the characteristic domains are found to involve a complex bifurcation structure, organized around three codimension-two bifurcation points: a Bogdanov-Takens point, a cusp point, and a fold-homoclinic point. Apart from the monostable domains, our study also reveals two domains admitting bistable behavior, manifested as coexistence between the two stationary solutions or between a stationary and a periodic solution. It is shown that the collective mode may emerge via two generic scenarios, guided by a saddle-node of infinite period or the Hopf bifurcation, such that the transition from the homogeneous to the heterogeneous stationary state under increasing diversity may follow the classical paradigm, but may also be hysteretic. We demonstrate that the basic bifurcation structure holds qualitatively in the presence of small noise or small coupling delay, with the boundaries of the characteristic domains shifted compared to the noiseless and the delay-free case.

Phase-sensitive excitability of a limit cycle.

The classical notion of excitability refers to an equilibrium state that shows under the influence of perturbations a nonlinear threshold-like behavior. Here, we extend this concept by demonstrating how periodic orbits can exhibit a specific form of excitable behavior where the nonlinear threshold-like response appears only after perturbations applied within a certain part of the periodic orbit, i.e., the excitability happens to be phase-sensitive. As a paradigmatic example of this concept, we employ the classical FitzHugh-Nagumo system. The relaxation oscillations, appearing in the oscillatory regime of this system, turn out to exhibit a phase-sensitive nonlinear threshold-like response to perturbations, which can be explained by the nonlinear behavior in the vicinity of the canard trajectory. Triggering the phase-sensitive excitability of the relaxation oscillations by noise, we find a characteristic non-monotone dependence of the mean spiking rate of the relaxation oscillation on the noise level. We explain this non-monotone dependence as a result of an interplay of two competing effects of the increasing noise: the growing efficiency of the excitation and the degradation of the nonlinear response.

Clustering promotes switching dynamics in networks of noisy neurons.

Macroscopic variability is an emergent property of neural networks, typically manifested in spontaneous switching between the episodes of elevated neuronal activity and the quiescent episodes. We investigate the conditions that facilitate switching dynamics, focusing on the interplay between the different sources of noise and heterogeneity of the network topology. We consider clustered networks of rate-based neurons subjected to external and intrinsic noise and derive an effective model where the network dynamics is described by a set of coupled second-order stochastic mean-field systems representing each of the clusters. The model provides an insight into the different contributions to effective macroscopic noise and qualitatively indicates the parameter domains where switching dynamics may occur. By analyzing the mean-field model in the thermodynamic limit, we demonstrate that clustering promotes multistability, which gives rise to switching dynamics in a considerably wider parameter region compared to the case of a non-clustered network with sparse random connection topology.

Mean-field dynamics of a population of stochastic map neurons.

We analyze the emergent regimes and the stimulus-response relationship of a population of noisy map neurons by means of a mean-field model, derived within the framework of cumulant approach complemented by the Gaussian closure hypothesis. It is demonstrated that the mean-field model can qualitatively account for stability and bifurcations of the exact system, capturing all the generic forms of collective behavior, including macroscopic excitability, subthreshold oscillations, periodic or chaotic spiking, and chaotic bursting dynamics. Apart from qualitative analogies, we find a substantial quantitative agreement between the exact and the approximate system, as reflected in matching of the parameter domains admitting the different dynamical regimes, as well as the characteristic properties of the associated time series. The effective model is further shown to reproduce with sufficient accuracy the phase response curves of the exact system and the assembly's response to external stimulation of finite amplitude and duration.

Phase response curves for models of earthquake fault dynamics.

We systematically study effects of external perturbations on models describing earthquake fault dynamics. The latter are based on the framework of the Burridge-Knopoff spring-block system, including the cases of a simple mono-block fault, as well as the paradigmatic complex faults made up of two identical or distinct blocks. The blocks exhibit relaxation oscillations, which are representative for the stick-slip behavior typical for earthquake dynamics. Our analysis is carried out by determining the phase response curves of first and second order. For a mono-block fault, we consider the impact of a single and two successive pulse perturbations, further demonstrating how the profile of phase response curves depends on the fault parameters. For a homogeneous two-block fault, our focus is on the scenario where each of the blocks is influenced by a single pulse, whereas for heterogeneous faults, we analyze how the response of the system depends on whether the stimulus is applied to the block having a shorter or a longer oscillation period.

Mean-field dynamics of a random neural network with noise.

We consider a network of randomly coupled rate-based neurons influenced by external and internal noise. We derive a second-order stochastic mean-field model for the network dynamics and use it to analyze the stability and bifurcations in the thermodynamic limit, as well as to study the fluctuations due to the finite-size effect. It is demonstrated that the two types of noise have substantially different impact on the network dynamics. While both sources of noise give rise to stochastic fluctuations in the case of the finite-size network, only the external noise affects the stationary activity levels of the network in the thermodynamic limit. We compare the theoretical predictions with the direct simulation results and show that they agree for large enough network sizes and for parameter domains sufficiently away from bifurcations.

Activation process in excitable systems with multiple noise sources: One and two interacting units.

We consider the coaction of two distinct noise sources on the activation process of a single excitable unit and two interacting excitable units, which are mathematically described by the Fitzhugh-Nagumo equations. We determine the most probable activation paths around which the corresponding stochastic trajectories are clustered. The key point lies in introducing appropriate boundary conditions that are relevant for a class II excitable unit, which can be immediately generalized also to scenarios involving two coupled units. We analyze the effects of the two noise sources on the statistical features of the activation process, in particular demonstrating how these are modified due to the linear or nonlinear form of interactions. Universal properties of the activation process are qualitatively discussed in the light of a stochastic bifurcation that underlies the transition from a stochastically stable fixed point to continuous oscillations.

Activation process in excitable systems with multiple noise sources: Large number of units.

We study the activation process in large assemblies of type II excitable units whose dynamics is influenced by two independent noise terms. The mean-field approach is applied to explicitly demonstrate that the assembly of excitable units can itself exhibit macroscopic excitable behavior. In order to facilitate the comparison between the excitable dynamics of a single unit and an assembly, we introduce three distinct formulations of the assembly activation event. Each formulation treats different aspects of the relevant phenomena, including the thresholdlike behavior and the role of coherence of individual spikes. Statistical properties of the assembly activation process, such as the mean time-to-first pulse and the associated coefficient of variation, are found to be qualitatively analogous for all three formulations, as well as to resemble the results for a single unit. These analogies are shown to derive from the fact that global variables undergo a stochastic bifurcation from the stochastically stable fixed point to continuous oscillations. Local activation processes are analyzed in the light of the competition between the noise-led and the relaxation-driven dynamics. We also briefly report on a system-size antiresonant effect displayed by the mean time-to-first pulse.

Persistence and failure of mean-field approximations adapted to a class of systems of delay-coupled excitable units.

We consider the approximations behind the typical mean-field model derived for a class of systems made up of type II excitable units influenced by noise and coupling delays. The formulation of the two approximations, referred to as the Gaussian and the quasi-independence approximation, as well as the fashion in which their validity is verified, are adapted to reflect the essential properties of the underlying system. It is demonstrated that the failure of the mean-field model associated with the breakdown of the quasi-independence approximation can be predicted by the noise-induced bistability in the dynamics of the mean-field system. As for the Gaussian approximation, its violation is related to the increase of noise intensity, but the actual condition for failure can be cast in qualitative, rather than quantitative terms. We also discuss how the fulfillment of the mean-field approximations affects the statistics of the first return times for the local and global variables, further exploring the link between the fulfillment of the quasi-independence approximation and certain forms of synchronization between the individual units.

Triggered dynamics in a model of different fault creep regimes.

The study is focused on the effect of transient external force induced by a passing seismic wave on fault motion in different creep regimes. Displacement along the fault is represented by the movement of a spring-block model, whereby the uniform and oscillatory motion correspond to the fault dynamics in post-seismic and inter-seismic creep regime, respectively. The effect of the external force is introduced as a change of block acceleration in the form of a sine wave scaled by an exponential pulse. Model dynamics is examined for variable parameters of the induced acceleration changes in reference to periodic oscillations of the unperturbed system above the supercritical Hopf bifurcation curve. The analysis indicates the occurrence of weak irregular oscillations if external force acts in the post-seismic creep regime. When fault motion is exposed to external force in the inter-seismic creep regime, one finds the transition to quasiperiodic- or chaos-like motion, which we attribute to the precursory creep regime and seismic motion, respectively. If the triggered acceleration changes are of longer duration, a reverse transition from inter-seismic to post-seismic creep regime is detected on a larger time scale.