The effects of time delay on the synchronization transitions in a modular neuronal network with hybrid synapses

Delay-induced synchronization transitions are studied in a modular neuronal network of small-world subnetworks with hybrid synapses in this paper. Numerical results show that the spatiotemporal synchronization transitions in a modular neuronal network not only depend on the information transmission delay, but also can be induced by the variations of the probability of inhibitory synapses and the number of subnetworks in the modular networks. In the hybrid modular network, the information transmission delay is shown to be significant, which can either promote or destroy synchronization of neuronal activity. In particular, the increasing delays can induce the intermittent appearance of regions of synchronization and non-synchronization. Interestingly, it is found that intermittent synchronization transition is relatively profound for smaller and larger probability of inhibitory synapses, while synchronization transition seems less profound for the moderate probability of inhibitory synapses. In addition, if only the delay is appropriate, there exists a suitable modular network topology structure enhancing the synchronized neuronal activity.     

Single or multiple synchronization transitions in scale-free neuronal networks with electrical or chemical coupling

In this paper, we have studied time delay- and coupling strength-induced synchronization transitions in scale-free modified Hodgkin–Huxley (MHH) neuron networks with gap-junctions and chemical synaptic coupling. It is shown that the synchronization transitions are much different for these two coupling types. For gap-junctions, the neurons exhibit a single synchronization transition with time delay and coupling strength, while for chemical synapses, there are multiple synchronization transitions with time delay, and the synchronization transition with coupling strength is dependent on the time delay lengths. For short delays we observe a single synchronization transition, whereas for long delays the neurons exhibit multiple synchronization transitions as the coupling strength is varied. These results show that gap junctions and chemical synapses have different impacts on the pattern formation and synchronization transitions of the scale-free MHH neuronal networks, and chemical synapses, compared to gap junctions, may play a dominant and more active function in the firing activity of the networks. These findings would be helpful for further understanding the roles of gap junctions and chemical synapses in the firing dynamics of neuronal networks.     

Stochastic resonance in small-world neuronal networks with hybrid electrical–chemical synapses

The dependence of stochastic resonance in small-world neuronal networks with hybrid electrical–chemical synapses on the probability of chemical synapse and the rewiring probability is investigated. A subthreshold periodic signal is imposed on one single neuron within the neuronal network as a pacemaker. It is shown that, irrespective of the probability of chemical synapse, there exists a moderate intensity of external noise optimizing the response of neuronal networks to the pacemaker. Moreover, the effect of pacemaker driven stochastic resonance of the system depends largely on the probability of chemical synapse. A high probability of chemical synapse will need lower noise intensity to evoke the phenomenon of stochastic resonance in the networked neuronal systems. In addition, for fixed noise intensity, there is an optimal chemical synapse probability, which can promote the propagation of the localized subthreshold pacemaker across neural networks. And the optimal chemical synapses probability turns even larger as the coupling strength decreases. Furthermore, the small-world topology has a significant impact on the stochastic resonance in hybrid neuronal networks. It is found that increasing the rewiring probability can always enhance the stochastic resonance until it approaches the random network limit.     

Ordering chaos and synchronization transitions by chemical delay and coupling on scale-free neuronal networks

Chemical synaptic connections are more common than electric ones in neurons, and information transmission delay is especially significant for the synapses of chemical type. In this paper, we report a phenomenon of ordering spatiotemporal chaos and synchronization transitions by the delays and coupling through chemical synapses of modified Hodgkin–Huxley (MHH) neurons on scale-free networks. As the delay τ is increased, the neurons exhibit transitions from bursting synchronization (BS) to intermittent multiple spiking synchronizations (SS). As the coupling g_syn is increased, the neurons exhibit different types of firing transitions, depending on the values of τ. For a smaller τ, there are transitions from spatiotemporal chaotic bursting (SCB) to BS or SS; while for a larger τ, there are transitions from SCB to intermittent multiple SS. These findings show that the delays and coupling through chemical synapses can tame the chaotic firings and repeatedly enhance the firing synchronization of neurons, and hence could play important roles in the firing activity of the neurons on scale-free networks.     

Spatiotemporal dynamics on small-world neuronal networks: The roles of two types of time-delayed coupling

We investigate temporal coherence and spatial synchronization on small-world networks consisting of noisy Terman–Wang (TW) excitable neurons in dependence on two types of time-delayed coupling: {x_j(t − τ) − x_i(t)} and {x_j(t − τ) − x_i(t − τ)}. For the former case, we show that time delay in the coupling can dramatically enhance temporal coherence and spatial synchrony of the noise-induced spike trains. In addition, if the delay time τ is tuned to nearly match the intrinsic spike period of the neuronal network, the system dynamics reaches a most ordered state, which is both periodic in time and nearly synchronized in space, demonstrating an interesting resonance phenomenon with delay. For the latter case, however, we cannot achieve a similar spatiotemporal ordered state, but the neuronal dynamics exhibits interesting synchronization transitions with time delay from zigzag fronts of excitations to dynamic clustering anti-phase synchronization (APS), and further to clustered chimera states which have spatially distributed anti-phase coherence separated by incoherence. Furthermore, we also show how these findings are influenced by the change of the noise intensity and the rewiring probability of the small-world networks. Finally, qualitative analysis is given to illustrate the numerical results.     

Optimal autaptic and synaptic delays enhanced synchronization transitions induced by each other in Newman–Watts neuronal networks

In this paper, we numerically study the effect of electrical autaptic and synaptic delays on synchronization transitions induced by each other in Newman–Watts Hodgkin–Huxley neuronal networks. It is found that the synchronization transitions induced by synaptic delay vary with varying autaptic delay and become strongest when autaptic delay is optimal. Similarly, the synchronization transitions induced by autaptic delay vary with varying synaptic delay and become strongest at optimal synaptic delay. Also, there is optimal coupling strength by which the synchronization transitions induced by either synaptic or autaptic delay become strongest. These results show that electrical autaptic and synaptic delays can enhance synchronization transitions induced by each other in the neuronal networks. This implies that electrical autaptic and synaptic delays can cooperate with each other and more efficiently regulate the synchrony state of the neuronal networks. These findings could find potential implications for the information transmission in neural systems.     

Impacts of hybrid synapses on the noise-delayed decay in scale-free neural networks

We study the phenomenon of noise-delayed decay in a scale-free neural network consisting of excitable FitzHugh–Nagumo neurons. In contrast to earlier works, where only electrical synapses are considered among neurons, we primarily examine the effects of hybrid synapses on the noise-delayed decay in this study. We show that the electrical synaptic coupling is more impressive than the chemical coupling in determining the appearance time of the first-spike and more efficient on the mitigation of the delay time in the detection of a suprathreshold input signal. We obtain that hybrid networks including inhibitory chemical synapses have higher signal detection capabilities than those of including excitatory ones. We also find that average degree exhibits two different effects, which are strengthening and weakening the noise-delayed decay effect depending on the noise intensity.     

The effects of synaptic time delay on motifs of chemically coupled Rulkov model neurons

Given the importance of the network motifs, we consider a pair of Rulkov chaotic map neurons, reciprocally coupled via symmetrical chemical synapses with the time delay τ. For the inhibitory and excitatory synapses, the system dynamics is determined by the synaptic weight g_c, synaptic gain parameter k, time delay τ and the external excitation σ. Due to chaotic nature of the map and synaptic model complexity, the appropriately averaged cross-correlation of membrane potentials represents a suitable numerical diagnostics to quantify mutual synchronization. Along with the expected phase and anti-phase synchronization regimes, we find the emergent phenomena that significantly influence the synchronization behavior.     

Adaptive synchronization in an array of chaotic neural networks with mixed delays and jumping stochastically hybrid coupling

In this paper, a general model of an array of N linearly coupled delayed neural networks with Markovian jumping hybrid coupling is introduced. The hybrid coupling consists of constant coupling, discrete and distributed time-varying delay coupling. The complex dynamical network jumps from one mode to another according to a Markovian chain, where all the coupling configurations are also dependent on mode switching. Meanwhile, all the coupling terms are subjected to stochastic disturbances which are described in terms of a Brownian motion. By adaptive approach, some sufficient criteria have been derived to ensure the synchronization in an array of jump neural networks with mixed delays and hybrid coupling in mean square. Surprisingly, it is found that complex networks with two different structure can also be synchronized according to known probability matrix. Finally, an example illustrated by switching between small-world networks and nearest-neighbor networks is given to show the effectiveness of the proposed criteria.     

Effects of time delay and coupling strength on synchronization transitions in excitable homogeneous random network

In this paper we numerically investigate the effects of time delay and coupling strength on synchronization transitions in excitable homogeneous random network. Different roles of time delay and coupling strength have been discovered by synchronization parameter and space–time plots. Specifically, we have found three distinct parameter regions, i.e., asynchronous region (domain I for small time delay), transition region (domain II for moderate time delay) and synchronous region (domain III for large time delay) as time delay is increased. The phenomenon of multi-stability is observed in the transition region. While coupling strength can enhance synchronization in the transition region and can reduce synchronization time in the synchronous region. All these results are independence on the system size.     

Delay-induced diversity of firing behavior and ordered chaotic firing in adaptive neuronal networks

In this paper, we study the effect of time delay on the firing behavior and temporal coherence and synchronization in Newman–Watts thermosensitive neuron networks with adaptive coupling. At beginning, the firing exhibit disordered spiking in absence of time delay. As time delay is increased, the neurons exhibit diversity of firing behaviors including bursting with multiple spikes in a burst, spiking, bursting with four, three and two spikes, firing death, and bursting with increasing amplitude. The spiking is the most ordered, exhibiting coherence resonance (CR)-like behavior, and the firing synchronization becomes enhanced with the increase of time delay. As growth rate of coupling strength or network randomness increases, CR-like behavior shifts to smaller time delay and the synchronization of firing increases. These results show that time delay can induce diversity of firing behaviors in adaptive neuronal networks, and can order the chaotic firing by enhancing and optimizing the temporal coherence and enhancing the synchronization of firing. However, the phenomenon of firing death shows that time delay may inhibit the firing of adaptive neuronal networks. These findings provide new insight into the role of time delay in the firing activity of adaptive neuronal networks, and can help to better understand the complex firing phenomena in neural networks.     

Multiple firing coherence resonances in excitatory and inhibitory coupled neurons

The impact of inhibitory and excitatory synapses in delay-coupled Hodgkin–Huxley neurons that are driven by noise is studied. If both synaptic types are used for coupling, appropriately tuned delays in the inhibition feedback induce multiple firing coherence resonances at sufficiently strong coupling strengths, thus giving rise to tongues of coherency in the corresponding delay-strength parameter plane. If only inhibitory synapses are used, however, appropriately tuned delays also give rise to multiresonant responses, yet the successive delays warranting an optimal coherence of excitations obey different relations with regards to the inherent time scales of neuronal dynamics. This leads to denser coherence resonance patterns in the delay-strength parameter plane. The robustness of these findings to the introduction of delay in the excitatory feedback, to noise, and to the number of coupled neurons is examined. Mechanisms underlying our observations are revealed, and it is suggested that the regularity of spiking across neuronal networks can be optimized in an unexpectedly rich variety of ways, depending on the type of coupling and the duration of delays.     

Endogenous fields enhanced stochastic resonance in a randomly coupled neuronal network

Endogenous field, evoked by structured neuronal network activity in vivo, is correlated with many vital neuronal processes. In this paper, the effects of endogenous fields on stochastic resonance (SR) in a randomly connected neuronal network are investigated. The network consists of excitatory and inhibitory neurons and the axonal conduction delays between neurons are also considered. Numerical results elucidate that endogenous field feedback results in more rhythmic macroscope activation of the network for proper time delay and feedback coefficient. The response of the network to the weak periodic stimulation can be notably enhanced by endogenous field feedback. Moreover, the endogenous field feedback delay plays a vital role in SR. We reveal that appropriately tuned delays of the feedback can either induce the enhancement of SR, appearing at every integer multiple of the weak input signal’s oscillation period, or the depression of SR, appearing at every integer multiple of half the weak input signal’s oscillation period for the same feedback coefficient. Interestingly, the parameters of low-passed filter which is used in obtaining the endogenous field feedback signal play a subtle role in SR.     

Functional motifs: a novel perspective on burst synchronization and regularization of neurons coupled via delayed inhibitory synapses

For one of the most common network motifs, an inhibitory neuron pair, we perform an extensive study of burst synchronization and the related phenomena applying the model of Rulkov maps coupled via delayed synapses. Instigated by the phase-plane analysis, that has the neuron switching between the noninteracting and the interacting map, it is demonstrated how the system evolution may be interpreted by means of the dynamical configurations of the motif, each represented by an extracted subgraph. Under the variation of the synaptic parameters, the probability of finding synchronized neurons in a given configuration is seen to reflect the way in which the anti-phase synchronization is eventually superseded by the synchronization in phase. Such an approach also provides a novel insight into regularization, characterizing the neuron bursting in either of these regimes. Looking into correlation of the two neurons’ bursting cycles we acquire a deeper understanding of the more sophisticated mechanisms by which the regularity in the time series is maintained. Further, it is examined whether introducing heterogeneity in the neuron or the synaptic parameters may prove advantageous over the homogeneous case with respect to burst synchronization.     

Vibrational resonance in neuron populations with hybrid synapses

In this paper, vibrational resonance in excitable neuron populations with synapses is investigated by numerical simulation. In particular, the effect of the hybrid synapses on the signal detection and transmission in neural system is studied. Different topologies from regular and random networks to small-world networks are considered to analyze the dependence of vibrational resonance on the network structure and parameters. It is shown that there exists an optimal amplitude of high-frequency driving, enhancing the response of coupled neuron populations to a subthreshold signal. We find that chemical synaptic coupling is more efficient than the electrical coupling in signal detection and electrical synaptic coupling is better in signal transmission. Neuron populations with hybrid synapses compromise the merits of the two types of coupling and have an advantage in information communication.     

Possibilities of introducing different functional circuits on top of a structural neuron triplet: Where do the gains lie?

Instigated by the research on clusterization phenomena in complex neural networks, we study a triplet of bursting Rulkov map neurons connected via inhibitory synapses with delay. It is demonstrated how on a background of structural motif one can build different types of functional circuits. The approach is based on utilizing the properties of the chemical synapses, whose gating is modeled by the fast threshold modulation, in conjunction with the phase plane analysis, allowing the system state to be represented in terms of maps the neurons reside on. For both the dynamical configurations, monitoring the layout of active neurons, and the functional motifs, following the maps where the synchronized neurons lie, we establish a one-on-one correspondence between sequences in the time series and the triads, making up the subgraphs of the original graph. By introducing the appropriate sets of quantities, one obtains not only the distributions of triads as a function of synaptic parameters, but is also able to identify a distinct triad whose presence may be viewed as a signature of the burst synchronization process. In another setup, the regularization of burst cycles for an arbitrary neuron is explained by classifying all the bursts as long or short, with their fractions linked to the abundances of triads under variation of synaptic parameters.     

Dependence of inter-neuronal effective connectivity on synchrony dynamics in neuronal network motifs

Effective connectivity, characterized as directional causal influences among neural units, is functionally significant to be reconstructed. Various dynamic regimes have been considered to underlie reshaping of the effective connections. In this work, the impact of zero-lag synchronization on the reconstruction of effective connectivity in neuronal network motifs is investigated. The synchronization analysis and effective connectivity estimation by using Granger causality (GC) method are performed. It is shown that the synchronization of the neurons at zero lag contributes to the reconstruction of reciprocal effective connections without synaptic connections. In addition, delay-induced zero-lag synchronous transition facilitates dynamic transformation of the causal interactions. With the increase of synaptic coupling strength, the causal interplay undergoes the transition to be statistically significant at a critical value. Furthermore, it can be found that multiple effective motifs are extracted from different synchronization states of the underlying structural motifs. GC measures of effective connectivity are proved to be reliable compared with the Information Flow for causal analysis. The obtained results may be helpful to future research about information processes.     

Bifurcation and synchronization of synaptically coupled FHN models with time delay

This paper presents an investigation of dynamics of the coupled nonidentical FHN models with synaptic connection, which can exhibit rich bifurcation behavior with variation of the coupling strength. With the time delay being introduced, the coupled neurons may display a transition from the original chaotic motions to periodic ones, which is accompanied by complex bifurcation scenario. At the same time, synchronization of the coupled neurons is studied in terms of their mean frequencies. We also find that the small time delay can induce new period windows with the coupling strength increasing. Moreover, it is found that synchronization of the coupled neurons can be achieved in some parameter ranges and related to their bifurcation transition. Bifurcation diagrams are obtained numerically or analytically from the mathematical model and the parameter regions of different behavior are clarified.     

Information processing,memories, and synchronization in chaotic neural network with the time delay

Information processing and two types of memory in an analog neural network model with time delay that produces chaos similar to the human and animal EEGs are considered. There are two levels of information processing in this neural network: the level of individual neurons and the level of the neural network. Similar to the state of brain, the state of chaotic neural network is defined. It is characterized by two types of memories (memory I and memory II) and correlation structure between the neurons. In normal (unperturbed) state, the neural network generates chaotic patterns of averaged neuronal activities (memory I) and patterns of oscillation amplitudes (memory II). In the presence of external stimulation, the activity patterns change, showing changes in both types of memory. As in experiments on stimulation of the brain, the neural network model shows synchronization of neuronal activities due to stimulus measured by Pearson's correlation coefficient. An increase in neural network asymmetry (increase of the neural network excitability) leads to the phenomenon similar to the epilepsy. Modeling of brain injury, Parkinson's disease, and dementia is performed by removing and weakening interneuron connections. In all cases, the chaotic neural network shows a decrease of the degree of chaos and changes in both types of memory similar to those observed in experiments with healthy human subjects and patients with Parkinson's disease and dementia. © 2005 Wiley Periodicals, Inc. Complexity 11:39–52, 2005     

Cooperative dynamics in neuronal networks

There exist rich cooperative behaviors and their transitions in biological neuronal systems as some key biological factors are changed. Among all of cooperative behaviors of neuronal systems, the existing experiments have shown that the spatiotemporal pattern and synchronization dynamics are very crucial, which are closely related to normal function and dysfunction of neuronal systems. Based on different neuron models, the recent works have been made to explore the mechanisms of pattern formation and synchronization transition. This paper mainly overviews the recent studies of the cooperative dynamics including the pattern formation and synchronization transition in biological neuronal networks. Firstly, we review complicated spatiotemporal pattern dynamics of neuronal networks. Secondly, the interesting synchronization transition is reviewed. Finally, conclusion is given and we put forward some outlooks of research on the cooperative behaviors in real neuronal networks.