In this paper, we focus on the emergence of diverse neuronal oscillations arising in a mixed population of neurons with different excitability properties. These properties produce mixed mode oscillations (MMOs) characterized by the combination of large amplitudes and alternate subthreshold or small amplitude oscillations. Considering the biophysically plausible, Izhikevich neuron model, we demonstrate that various MMOs, including MMBOs (mixed mode bursting oscillations) and synchronized tonic spiking appear in a randomly connected network of neurons, where a fraction of them is in a quiescent (silent) state and the rest in self-oscillatory (firing) states. We show that MMOs and other patterns of neural activity depend on the number of oscillatory neighbors of quiescent nodes and on electrical coupling strengths. Our results are verified by constructing a reduced-order network model and supported by systematic bifurcation diagrams as well as for a small-world network. Our results suggest that, for weak couplings, MMOs appear due to the de-synchronization of a large number of quiescent neurons in the networks. The quiescent neurons together with the firing neurons produce high frequency oscillations and bursting activity. The overarching goal is to uncover a favorable network architecture and suitable parameter spaces where Izhikevich model neurons generate diverse responses ranging from MMOs to tonic spiking.
In this work, we study the dynamic range of a neuronal network of excitable neurons with excitatory and inhibitory synapses. We obtain an analytical expression for the critical point as a function of the excitatory and inhibitory synaptic intensities. We also determine an analytical expression that gives the critical point value in which the maximal dynamic range occurs. Depending on the mean connection degree and coupl\-ing weights, the critical points can exhibit ceasing or ceaseless dynamics. However, the dynamic range is equal in both cases. We observe that the external stimulus mask some …
We describe a form of memory exhibited by extended excitable systems driven by stochastic fluctuations. Under such conditions, the system self-organizes into a state characterized by power-law correlations thus retaining long-term memory of previous states. The exponents are robust and model-independent. We discuss novel implications of these results for the functioning of cortical neurons as well as for networks of neurons.
The study of balanced networks of excitatory and inhibitory neurons has led to several open questions. On the one hand it is yet unclear whether the asynchronous state observed in the brain is autonomously generated, or if it results from the interplay between external drivings and internal dynamics. It is also not known, which kind of network variabilities will lead to irregular spiking and which to synchronous firing states. Here we show how isolated networks of purely excitatory neurons generically show asynchronous firing whenever a minimal level of structural variability is present togeth…
Spontaneous formation of clusters of synchronized spiking in a structureless ensemble of equal stochastically perturbed excitable neurons with delayed coupling is demonstrated for the first time. The effect is a consequence of a subtle interplay between interaction delays, noise and the excitable character of a single neuron. Dependence of the cluster properties on the time-lag, noise intensity and the synaptic strength is investigated.
We demonstrate that chimera behavior can be observed in nonlocally coupled networks of excitable systems in the presence of noise. This phenomenon is distinct from classical chimeras, which occur in deterministic oscillatory systems, and it combines temporal features of coherence resonance, i.e., the constructive role of noise, and spatial properties of chimera states, i.e., coexistence of spatially coherent and incoherent domains in a network of identical elements. Coherence-resonance chimeras are associated with alternating switching of the location of coherent and incoherent domains, which …
We study the emergence and coherence of stochastic oscillations in star networks of excitable elements in which peripheral nodes receive independent random inputs. A biophysical model of a distal branch of sensory neuron in which peripheral nodes of Ranvier are coupled to a central node by myelinated cable segments is used along with a generic model of networked stochastic active rotators. We show that coherent oscillations can emerge due to stochastic synchronization of peripheral nodes and that the degree of coherence can be maximized by tuning the coupling strength and the size of the netwo…
We study the collective dynamics in a network of excitable units (neurons) adaptively interacting with a pool of resources. The resource pool is influenced by the average activity of the network, whereas the feedback from the resources to the network is comprised of components acting homogeneously or inhomogeneously on individual units of the network. 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 network and the resources can give rise to collective activity bursting in the network. To explain the…
We study the stochastic dynamics of strongly-coupled excitable elements on a tree network. The peripheral nodes receive independent random inputs which may induce large spiking events propagating through the branches of the tree and leading to global coherent oscillations in the network. This scenario may be relevant to action potential generation in certain sensory neurons, which possess myelinated distal dendritic tree-like arbors with excitable nodes of Ranvier at peripheral and branching nodes and exhibit noisy periodic sequences of action potentials. We focus on the spiking statistics of …
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…
We derive the Kuramoto model (KM) corresponding to a population of weakly coupled, nearly identical quadratic integrate-and-fire (QIF) neurons with both electrical and chemical coupling. The ratio of chemical to electrical coupling determines the phase lag of the characteristic sine coupling function of the KM and critically determines the synchronization properties of the network. We apply our results to investigate chimera states in two coupled populations of identical QIF neurons. We find that the presence of both electrical and chemical coupling is a necessary condition for chimera states …
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 busting. 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 as a paradigmatic model for a neuron with a slow recovery variable or, more generally, as an excitable system under the influe…
We study rotating waves in the Theta model for a ring of synaptically-interacting neurons. We prove that when the neurons are oscillatory, at least one rotating wave always exists. In the case of excitable neurons, we prove that no travelling waves exist when the synaptic coupling is weak, and at least two rotating waves, a `fast' one and a `slow' one, exist when the synaptic coupling is sufficiently strong. We derive explicit upper and lower bounds for the `critical' coupling strength as well as for wave velocities. We also study the special case of uniform coupling, in which complete analyti…
Chemical and electrical synapses shape the dynamics of neuronal networks. Numerous theoretical studies have investigated how each of these types of synapses contributes to the generation of neuronal oscillations, but their combined effect is less understood. This limitation is further magnified by the impossibility of traditional neuronal mean-field models ---also known as firing rate models, or firing rate equations--- to account for electrical synapses. Here we introduce a novel firing rate model that exactly describes the mean field dynamics of heterogeneous populations of quadratic integra…
Electrical coupling between neurons is broadly present across brain areas and is typically assumed to synchronize network activity. However, intrinsic properties of the coupled cells can complicate this simple picture. Many cell types with strong electrical coupling have been shown to exhibit resonant properties, and the subthreshold fluctuations arising from resonance are transmitted through electrical synapses in addition to action potentials. Using the theory of weakly coupled oscillators, we explore the effect of both subthreshold and spike-mediated coupling on synchrony in small networks …
We study periodic travelling waves in the Theta model for a linear continuum of synaptically-interacting neurons. We prove that when the neurons are oscillatory, at least one periodic travelling of every wave number always exists. In the case of excitable neurons, we prove that no periodic travelling waves exist when the synaptic coupling is weak, and at least two periodic travelling waves of each wave-number, a `fast' one and a `slow' one, exist when the synaptic coupling is sufficiently strong. We derive explicit upper and lower bounds for the `critical' coupling strength as well as for the …
Experiments suggest that cerebral cortex gains several functional advantages by operating in a dynamical regime near the critical point of a phase transition. However, a long-standing criticism of this hypothesis is that critical dynamics are rather noisy, which might be detrimental to aspects of brain function that require precision. If the cortex does operate near criticality, how might it mitigate the noisy fluctuations? One possibility is that other parts of the brain may act to control the fluctuations and reduce cortical noise. To better understand this possibility, here we numerically a…
We study the macroscopic dynamics of large networks of excitable type 1 neurons composed of two populations interacting with disparate but symmetric intra- and inter-population coupling strengths. This nonuniform coupling scheme facilitates symmetric equilibria, where both populations display identical firing activity, characterized by either quiescent or spiking behavior, or asymmetric equilibria, where the firing activity of one population exhibits quiescent but the other exhibits spiking behavior. Oscillations in the firing rate are possible if neurons emit pulses with non-zero width but ar…
We study synaptically coupled neuronal networks to identify the role of coupling delays in network's synchronized behaviors. We consider a network of excitable, relaxation oscillator neurons where two distinct populations, one excitatory and one inhibitory, are coupled and interact with each other. The excitatory population is uncoupled, while the inhibitory population is tightly coupled. A geometric singular perturbation analysis yields existence and stability conditions for synchronization states under different firing patterns between the two populations, along with formulas for the periods…
Electrical synapses play a major role in setting up neuronal synchronization, but the precise mechanisms whereby these synapses contribute to synchrony are subtle and remain elusive. To investigate these mechanisms mean-field theories for quadratic integrate-and-fire neurons with electrical synapses have been recently put forward. Still, the validity of these theories is controversial since they assume that the neurons produce unrealistic, symmetric spikes, ignoring the well-known impact of spike shape on synchronization. Here we show that the assumption of symmetric spikes can be relaxed in s…
Many experiments have evidenced that electrical and chemical synapses -- hybrid synapses -- coexist in most organisms and brain structures. The role of electrical and chemical synapse connection in diversity of neural activity generation has been investigated separately in networks of varying complexities. Nevertheless, theoretical understanding of hybrid synapses in diverse dynamical states of neural networks for self-organization and robustness still has not been fully studied. Here, we present a model of neural network built with hybrid synapses to investigate the emergence of global and co…
Here we study the emergence of chimera states, a recently reported phenomenon referring to the coexistence of synchronized and unsynchronized dynamical units, in a population of Morris-Lecar neurons which are coupled by both electrical and chemical synapses, constituting a hybrid synaptic architecture, as in actual brain connectivity. This scheme consists of a nonlocal network where the nearest neighbor neurons are coupled by electrical synapses, while the synapses from more distant neurons are of the chemical type. We demonstrate that peculiar dynamical behaviors, including chimera state and …
We consider large networks of globally coupled spiking neurons and derive an exact low-dimensional description of their collective dynamics in the thermodynamic limit. Individual neurons are described by the Ermentrout-Kopell canonical model that can be excitable or tonically spiking, and interact with other neurons via pulses. Utilizing the equivalence of the quadratic integrate- and-fire and the theta neuron formulations, we first derive the dynamical equations in terms of the Kuramoto-Daido order parameters (Fourier modes of the phase distribution) and relate them to two biophysically relev…
The phenomenon of self-induced stochastic resonance (SISR) requires a nontrivial scaling limit between the deterministic and the stochastic timescales of an excitable system, leading to the emergence of coherent oscillations which are absent without noise. In this paper, we numerically investigate SISR and its control in single neurons and three-neuron motifs made up of the Morris-Lecar model. In single neurons, we compare the effects of electrical and chemical autapses on the degree of coherence of the oscillations due to SISR. In the motifs, we compare the effects of altering the synaptic ti…
We study the dynamic range of a cellular automaton model for a neuronal network with electrical and chemical synapses. The neural network is separated into two layers, where one layer corresponds to inhibitory, and the other corresponds to excitatory neurons. We randomly distribute electrical synapses in the network, in order to analyse the effects on the dynamic range. We verify that electrical synapses have a complementary effect on the enhancement of the dynamic range. The enhancement depends on the proportion of electrical synapses, and also depends on the layer where they are distributed.
Recurrent networks of dynamic elements frequently exhibit emergent collective oscillations, which can display substantial regularity even when the individual elements are considerably noisy. How noise-induced dynamics at the local level coexists with regular oscillations at the global level is still unclear. Here we show that a combination of stochastic recurrence-based initiation with deterministic refractoriness in an excitable network can reconcile these two features, leading to maximum collective coherence for an intermediate noise level. We report this behavior in the slow oscillation reg…
In the last decade, several models with network adaptive mechanisms (link deletion-creation, dynamic synapses, dynamic gains) have been proposed as examples of self-organized criticality (SOC) to explain neuronal avalanches. However, all these systems present stochastic oscillations hovering around the critical region that are incompatible with standard SOC. This phenomenology has been called self-organized quasi-criticality (SOqC). Here we make a linear stability analysis of the mean field fixed points of two SOqC systems: a fully connected network of discrete time stochastic spiking neurons …
It has been found that contrarian oscillators usually take a negative role in the collective behaviors formed by conformist oscillators. However, experiments revealed that it is also possible to achieve a strong coherence even when there are contrarians in the system such as neuron networks with both excitable and inhibitory neurons. To understand the underlying mechanism of this abnormal phenomenon, we here consider a complex network of coupled Kuramoto oscillators with mixed positive and negative couplings and present an efficient approach, i.e. tit-for-tat strategy, to suppress the negative…
Electrical stimulation of neural systems is a key tool for understanding neural dynamics and ultimately for developing clinical treatments. Many applications of electrical stimulation affect large populations of neurons. However, computational models of large networks of spiking neurons are inherently hard to simulate and analyze. We evaluate a reduced mean-field model of excitatory and inhibitory adaptive exponential integrate-and-fire (AdEx) neurons which can be used to efficiently study the effects of electrical stimulation on large neural populations. The rich dynamical properties of this …
