小世界神经元网络随机共振现象:混合突触和部分时滞的影响

实际神经元网络中,信息传递时电突触和化学突触同时存在,并且有些神经元间的时滞很小可以忽略.本文构建了带有不同类型突触耦合的小世界网络,研究部分时滞、混合突触及噪声对随机共振的影响.结果表明:兴奋性和抑制性突触的比例影响共振的产生;在抑制性突触为主的网络里,几乎不产生随机共振.系统最佳噪声强度和化学突触比例大致呈线性递增关系;特别是在以化学耦合为主的混合突触网络里,仅当兴奋性突触与抑制性突触比例约为4:1时,噪声才可诱导网络产生共振行为.在此比例下,引入部分时滞发现时滞可诱导网络产生随机多共振,且随网络中时滞边比例的增加,系统响应强度达到最优水平的时滞取值区间逐渐变窄;同时发现,网络中含有的化学突触越多,部分时滞诱导产生的多共振行为越强.此外,当时滞为系统固有周期的整数倍时,时滞越大共振所对应的噪声区域越广;并且网络中时滞边越多,越容易促使噪声和时滞诱导其产生明显的共振行为.     

Delay-induced synchronization transitions in small-world neuronal networks with hybrid electrical and chemical synapses

We study the dependence of synchronization transitions in small-world networks of bursting neurons with hybrid electrical–chemical synapses on the information transmission delay, the probability of electrical synapses, and the rewiring probability. It is shown that, irrespective of the probability of electrical synapses, the information transmission delay can always induce synchronization transitions in small-world neuronal networks, i.e., regions of synchronization and nonsynchronization appear intermittently as the delay increases. In particular, all these transitions to burst synchronization occur approximately at integer multiples of the bursting period of individual neurons. In addition, for larger probability of electrical synapses, the intermittent synchronization transition is more profound, due to the stronger synchronization ability of electrical synapses compared with chemical ones. More importantly, chemical and electrical synapses can perform complementary roles in the synchronization of hybrid small-world neuronal networks: the larger the electrical synapse strength is, the smaller the chemical synapse strength needed to achieve burst synchronization. Furthermore, the small-world topology has a significant effect on the synchronization transition in hybrid neuronal networks. It is found that increasing the rewiring probability can always enhance the synchronization of neuronal activity. The results obtained are instructive for understanding the synchronous behavior of neural systems.     

Impact of delays and rewiring on the dynamics of small-world neuronal networks with two types of coupling

We study synchronization transitions and pattern formation on small-world networks consisting of Morris-Lecar excitable neurons in dependence on the information transmission delay and the rewiring probability. In addition, networks formed via gap junctional connections and coupling via chemical synapses are considered separately. For gap-junctionally coupled networks we show that short delays can induce zigzag fronts of excitations, whereas long delays can further detriment synchronization due to a dynamic clustering anti-phase synchronization transition. For the synaptically coupled networks, on the other hand, we find that the clustering anti-phase synchronization can appear as a direct consequence of the prolongation of information transmission delay, without being accompanied by zigzag excitatory fronts. Irrespective of the coupling type, however, we show that an appropriate small-world topology can always restore synchronized activity if only the information transmission delays are short or moderate at most. Long information transmission delays always evoke anti-phase synchronization and clustering, in which case the fine-tuning of the network topology fails to restore the synchronization of neuronal activity.     

In- and Anti-transition of Firing Patterns Induced by Random Long-range Connections in Coupled Hindmarsh-Rose Neurons System

研究了在外界刺激电流的作用下,随机的长程关联对耦合的Hindmarsh-Rose神经元放电模式转变的影响．结果表明,当耦合强度较弱时,在神经元网络中加入一定数量的随机的长程关联,神经元的放电模式会从较少的周期态转变到较多的周期态;当耦合强度较强时,在神经元网络中加入一定数量的随机长程关联,神经元的放电模式会产生相反的转变,即从较多的周期态转变到较少的周期态．同时还简单讨论了神经系统的尺度大小和神经元之间的耦合强度,以及不同外界刺激条件下放电模式的强度与临界特性之间的关系．     

动态突触、神经耦合与时间延迟对神经元发放的影响

神经元膜电位的受激发放在神经系统的信息传递中起着重要作用.基于一个受动态突触刺激的突触后神经元发放模型,采用数值模拟和傅里叶变换分析的方法研究了动态突触、神经耦合与时间延迟对突触后神经元发放的影响.结果发现:突触前神经元发放频率与Hodgkin-Huxley神经元的固有频率发生共振决定了突触后神经元发放的难易,特定频率范围内的电流刺激有利于神经元激发,动态突触输出的随机突触电流中这些电流刺激所占的比率在很大程度上影响了突触后神经元的发放次数;将突触后神经元换成神经网络后,网络中神经元之间的耦合可以促进神经元的发放,耦合中的时间延迟可以增强这种促进作用,但是不会改变神经耦合对神经元发放的促进模式.     

Multiple attractors, long chaotic transients, and failure in small-world networks of excitable neurons

We study the dynamical states of a small-world network of recurrently coupled excitable neurons, through both numerical and analytical methods. The dynamics of this system depend mostly on both the number of long-range connections or "shortcuts", and the delay associated with neuronal interactions. We find that persistent activity emerges at low density of shortcuts, and that the system undergoes a transition to failure as their density reaches a critical value. The state of persistent activity below this transition consists of multiple stable periodic attractors, whose number increases at least as fast as the number of neurons in the network. At large shortcut density and for long enough delays the network dynamics exhibit exceedingly long chaotic transients, whose failure times follow a stretched exponential distribution. We show that this functional form arises for the ensemble-averaged activity if the failure time for each individual network realization is exponentially distributed.     

部分时滞诱发Watts-Strogatz小世界神经元网络产生随机多共振

已有研究显示时滞可诱发神经元网络产生随机多共振,但它们主要讨论了神经元间的耦合都存在时滞的情形.然而实际中,有些神经元间的信息传递是瞬时的或时滞很小可以忽略的,即神经元网络中只有部分神经元间的耦合具有时滞,简称部分时滞(若神经元网络内共有l条耦合边,其中有l1条耦合边是具有时滞的,而剩余的耦合边的时滞为零,则我们称这类时滞为部分时滞).本文以Watts-Strogatz小世界神经元网络为研究对象,主要讨论部分时滞对该神经元网络系统响应强度的影响.研究结果指出,系统响应强度随部分时滞的增加呈现多峰的变化态势,即部分时滞可诱发随机多共振现象;而且使系统响应强度达到最优水平的部分时滞的取值区间随随机时滞边概率的增加渐渐变窄,当随机时滞边概率足够大时,系统响应强度只有在时滞位于外界信号周期的整数倍附近才会达到最优.此外,我们还分析了随机连边概率和神经元网络中边的总数对部分时滞诱发的随机多共振现象的影响.结果显示,部分时滞诱发的随机多共振现象对随机连边概率具有一定的鲁棒性,而神经元网络中边的总数对部分时滞诱发的随机多共振的影响则较大.     

Phase synchronization of bursting neural networks with electrical and delayed dynamic chemical couplings

Diffusive electrical connections in neuronal networks are instantaneous, while excitatoryor inhibitory couplings through chemical synapses contain a transmission time-delay.Moreover, chemical synapses are nonlinear dynamical systems whose behavior can bedescribed by nonlinear differential equations. In this work, neuronal networks withdiffusive electrical couplings and time-delayed dynamic chemical couplings are considered.We investigate the effects of distributed time delays on phase synchronization of burstingneurons. We observe that in both excitatory and Inhibitory chemical connections, the phasesynchronization might be enhanced when time-delay is taken into account. This distributedtime delay can induce a variety of phase-coherent dynamical behaviors. We also study thecollective dynamics of network of bursting neurons. The network model presents theso-called Small-World property, encompassing neurons whose dynamics have two time scales(fast and slow time scales). The neuron parameters in such Small-World network, aresupposed to be slightly different such that, there may be synchronization of the bursting(slow) activity if the coupling strengths are large enough. Bounds for the criticalcoupling strengths to obtain burst synchronization in terms of the network structure aregiven. Our studies show that the network synchronizability is improved, as itsheterogeneity is reduced. The roles of synaptic parameters, more precisely those of thecoupling strengths and the network size are also investigated.     

Stochastic resonance in hybrid scale-free neuronal networks

We study the phenomenon of stochastic resonance in a system of coupled neurons that are globally excited by a weak periodic input signal. We make the realistic assumption that the chemical and electrical synapses interact in the same neuronal network, hence constituting a hybrid network. By considering a hybrid coupling scheme embedded in the scale-free topology, we show that the electrical synapses are more efficient than chemical synapses in promoting the best correlation between the weak input signal and the response of the system. We also demonstrate that the average degree of neurons within the hybrid scale-free network significantly influences the optimal amount of noise for the occurrence of stochastic resonance, indicating that there also exists an optimal topology for the amplification of the response to the weak input signal. Lastly, we verify that the presented results are robust to variations of the system size.     

Effects of the network structure and coupling strength on the noise-induced response delay of a neuronal network

The Hodgkin-Huxley (H-H) neuron model driven by stimuli just above threshold shows a noise-induced response delay with respect to time to the first spike for a certain range of noise strengths, an effect called “noise delayed decay” (NDD). We study the response time of a network of coupled H-H neurons, and investigate how the NDD can be affected by the connection topology of the network and the coupling strength. We show that the NDD effect exists for weak and intermediate coupling strengths, whereas it disappears for strong coupling strength regardless of the connection topology. We also show that although the network structure has very little effect on the NDD for a weak coupling strength, the network structure plays a key role for an intermediate coupling strength by decreasing the NDD effect with the increasing number of random shortcuts, and thus provides an additional operating regime, that is absent in the regular network, in which the neurons may also exploit a spike time code.     

Inferring Excitatory and Inhibitory Connections in Neuronal Networks

The comprehension of neuronal network functioning, from most basic mechanisms of signal transmission to complex patterns of memory and decision making, is at the basis of the modern research in experimental and computational neurophysiology. While mechanistic knowledge of neurons and synapses structure increased, the study of functional and effective networks is more complex, involving emergent phenomena, nonlinear responses, collective waves, correlation and causal interactions. Refined data analysis may help in inferring functional/effective interactions and connectivity from neuronal activity. The Transfer Entropy (TE) technique is, among other things, well suited to predict structural interactions between neurons, and to infer both effective and structural connectivity in small- and large-scale networks. To efficiently disentangle the excitatory and inhibitory neural activities, in the article we present a revised version of TE, split in two contributions and characterized by a suited delay time. The method is tested on in silico small neuronal networks, built to simulate the calcium activity as measured via calcium imaging in two-dimensional neuronal cultures. The inhibitory connections are well characterized, still preserving a high accuracy for excitatory connections prediction. The method could be applied to study effective and structural interactions in systems of excitable cells, both in physiological and in pathological conditions.     

Firing activity of complex space-clamped FitzHugh-Nagumo neural networks

We investigate how firing activity of complex neural networks depends on the random long-range connections and coupling strength. Network elements are described by excitable space-clamped FitzHugh-Nagumo (SCFHN) neurons with the values of parameters at which no firing activity occurs. It is found that for a given appropriate coupling strength C, there exists a critical fraction of random connections (or randomness) p^*, such that if p > p^* the firing neurons, which are absent in the nearest-neighbor network, occur. The firing activity becomes more frequent as randomness p is further increased. On the other hand, when the p is smaller, there are no active neurons in network, no matter what the value of C is. For a given larger p, there exist optimal coupling strength levels, where firing activity reaches its maximum. To the best of our knowledge, this is a novel mechanism for the emergence of firing activity in neurons.     

Topological probability and connection strength induced activity in complex neural networks

Recent experimental evidence suggests that some brain activities can be assigned to small-world networks. In this work, we investigate how the topological probability p and connection strength C affect the activities of discrete neural networks with small-world (SW) connections. Network elements are described by two-dimensional map neurons (2DMNs) with the values of parameters at which no activity occurs. It is found that when the value of p is smaller or larger, there are no active neurons in the network, no matter what the value of connection strength is; for a given appropriate connection strength, there is an intermediate range of topological probability where the activity of 2DMN network is induced and enhanced. On the other hand, for a given intermediate topological probability level, there exists an optimal value of connection strength such that the frequency of activity reaches its maximum. The possible mechanism behind the action of topological probability and connection strength is addressed based on the bifurcation method. Furthermore, the effects of noise and transmission delay on the activity of neural network are also studied.     

Synchronization scenarios of chimeras in multiplex networks

Complex networks consisting of several interacting layers allow for remote synchronization of distant layers via an intermediate relay layer. We extend the notion of relay synchronization to chimera states, and study the scenarios of relay synchronization in a three-layer network of FitzHugh–Nagumo (FHN) oscillators, where each layer has a nonlocal coupling topology. Varying the coupling strength and time delay in the inter-layer connections, we observe relay synchronization between chimera states, i.e., complex spatio-temporal patterns of coexisting coherent and incoherent domains, in the outer network layers. Special regimes where only the coherent domains of chimeras are synchronized, and the incoherent domains remain desynchronized, as well as transitions between different synchronization regimes are analyzed.     

Model architecture for associative memory in a neural network of spiking neurons

A synaptic connectivity model is assembled on a spiking neuron network aiming to build up a dynamic pattern recognition system. The connection architecture includes gap junctions and both inhibitory and excitatory chemical synapses based on Hebb’s hypothesis. The network evolution resulting from external stimulus is sampled in a properly defined frequency space. Neurons’ responses to different current injections are mapped onto a subspace using Principal Component Analysis. Departing from the base attractor, related to a quiescent state, different external stimuli drive the network to different fixed points through specific trajectories in this subspace.     

Small-World Connections to Induce Firing Activity and Phase Synchronization in Neural Networks

We investigate how the firing activity and the subsequent phase synchronization of neural networks with smallworld topological connections depend on the probability p of adding-links. Network elements are described by two-dimensional map neurons （2DMNs） in a quiescent original state. Neurons burst for a given coupling strength when the topological randomness p increases, which is absent in a regular-lattice neural network. The bursting activity becomes frequent and synchronization of neurons emerges as topological randomness further increases. The maximal firing frequency and phase synchronization appear at a particular value of p. However, if the randomness p further increases, the firing frequency decreases and synchronization is apparently destroyed.     

Delay-induced intermittent transition of synchronization in neuronal networks with hybrid synapses

We study the dependence of synchronization transitions in scale-free networks of bursting neurons with hybrid synapses on the information transmission delay and the probability of inhibitory synapses. It is shown that, irrespective of the probability of inhibitory synapses, the delay always plays a subtle role during synchronization transition of the scale-free neuronal networks. In particular, regions of irregular and regular propagating excitatory fronts appear intermittently as the delay increases. These delay-induced synchronization transitions are manifested as well-expressed minima in the measure for spatiotemporal synchrony. In addition, it is found that, for smaller and larger probability of inhibitory synapses, intermittent synchronization transition is relatively profound, while for the moderate probability of inhibitory synapses, synchronization transition seems less profound. More interestingly, it is found that as the probability of inhibitory synapses is large, regions of synchronization are upscattering.     

The weakly nonlocal limit of a one-population Wilson-Cowan model

We derive the weakly nonlocal limit of a one-population neuronal field model of the Wilson-Cowan type in one spatial dimension. By transforming this equation to an equation in the firing rate variable, it is shown that stationary periodic solutions exist by appealing to a pseudo-potential analysis. The solutions of the full nonlocal equation obey a uniform bound, and the stationary periodic solutions in the weakly nonlocal limit satisfying the same uniform bound are characterized by finite ranges of pseudo energy constants. The time dependent version of the model is reformulated as a Ginzburg-Landau-Khalatnikov type of equation in the firing rate variable where the maximum (minimum) points correspond stable (unstable) homogeneous solutions of the weakly nonlocal limit. Based on this formulation it is also conjectured that the stationary periodic solutions are unstable. We implement a numerical method for the weakly nonlocal limit of the Wilson-Cowan type of model based on the wavelet-Galerkin approach. We perform some numerical tests to illustrate the stability of homogeneous solutions and the evolution of the bumps.     

Shift of percolation thresholds for epidemic spread between static and dynamic small-world networks

The study compares the epidemic spread on static and dynamic small-world networks. They are constructed as a 2-dimensional Newman and Watts model (500 × 500 square lattice with additional shortcuts), where the dynamics involves rewiring shortcuts in every time step of the epidemic spread. We assume susceptible-infectious-removed (SIR) model of the disease. We study the behaviour of the epidemic over the range of shortcut probability per underlying bond ϕ = 0–0.5. We calculate percolation thresholds for the epidemic outbreak, for which numerical results are checked against an approximate analytical model. We find a significant lowering of percolation thresholds on the dynamic network in the parameter range given. The result shows the behaviour of the epidemic on dynamic network is that of a static small world with the number of shortcuts increased by 20.7±1.4 %, while the overall qualitative behaviour stays the same. We derive corrections to the analytical model which account for the effect. For both dynamic and static small worlds we observe suppression of the average epidemic size dependence on network size in comparison with the finite-size scaling known for regular lattice. We also study the effect of dynamics for several rewiring rates relative to infectious period of the disease.