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.     

场耦合忆阻神经网络的电活动

神经元之间除了突触耦合,还存在磁通耦合.因此在传统的神经元模型中引入磁通量,并研究场耦合下神经网络的放电活动具有实际意义.建立一个含场耦合的Hodgkin-Huxley忆阻神经网络,引入神经元节点之间的距离权重,用磁通量描述时变电磁场,采用磁控忆阻器实现膜电位和磁通量之间的耦合.探讨距离权重和系统大小对神经网络放电活动的影响.研究发现,随着权重增大,神经网络放电活动增强,且系统规模越大,诱导神经元兴奋性的权重阈值越大,系统大小不影响神经网络活性随距离权重变化的规律.在不同的权重值下,神经网络活性随系统大小变化的规律明显不同.研究表明,距离权重和系统大小对含场耦合的忆阻神经网络放电活动有重要影响,其中距离权重起主导作用.     

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.     

Vibrational resonance in excitable neuronal systems

In this paper, we investigate the effect of a high-frequency driving on the dynamical response of excitable neuronal systems to a subthreshold low-frequency signal by numerical simulation. We demonstrate the occurrence of vibrational resonance in spatially extended neuronal networks. Different network topologies from single small-world networks to modular networks of small-world subnetworks are considered. It is shown that an optimal amplitude of high-frequency driving enhances the response of neuron populations to a low-frequency signal. This effect of vibrational resonance of neuronal systems depends extensively on the network structure and parameters, such as the coupling strength between neurons, network size, and rewiring probability of single small-world networks, as well as the number of links between different subnetworks and the number of subnetworks in the modular networks. All these parameters play a key role in determining the ability of the network to enhance the outreach of the localized subthreshold low-frequency signal. Considering that two-frequency signals are ubiquity in brain dynamics, we expect the presented results could have important implications for the weak signal detection and information propagation across neuronal systems.     

Automatic Search Dense Connection Module for Super-Resolution

The development of display technology has continuously increased the requirements for image resolution. However, the imaging systems of many cameras are limited by their physical conditions, and the image resolution is often restrictive. Recently, several models based on deep convolutional neural network (CNN) have gained significant performance for image super-resolution (SR), while extensive memory consumption and computation overhead hinder practical applications. For this purpose, we present a lightweight network that automatically searches dense connection (ASDCN) for image super-resolution (SR), which effectively reduces redundancy in dense connection and focuses on more valuable features. We employ neural architecture search (NAS) to model the searching of dense connections. Qualitative and quantitative experiments on five public datasets show that our derived model achieves superior performance over the state-of-the-art models.     

On the Nature of Functional Differentiation: The Role of Self-Organization with Constraints

The focus of this article is the self-organization of neural systems under constraints. In 2016, we proposed a theory for self-organization with constraints to clarify the neural mechanism of functional differentiation. As a typical application of the theory, we developed evolutionary reservoir computers that exhibit functional differentiation of neurons. Regarding the self-organized structure of neural systems, Warren McCulloch described the neural networks of the brain as being “heterarchical”, rather than hierarchical, in structure. Unlike the fixed boundary conditions in conventional self-organization theory, where stationary phenomena are the target for study, the neural networks of the brain change their functional structure via synaptic learning and neural differentiation to exhibit specific functions, thereby adapting to nonstationary environmental changes. Thus, the neural network structure is altered dynamically among possible network structures. We refer to such changes as a dynamic heterarchy. Through the dynamic changes of the network structure under constraints, such as physical, chemical, and informational factors, which act on the whole system, neural systems realize functional differentiation or functional parcellation. Based on the computation results of our model for functional differentiation, we propose hypotheses on the neuronal mechanism of functional differentiation. Finally, using the Kolmogorov–Arnold–Sprecher superposition theorem, which can be realized by a layered deep neural network, we propose a possible scenario of functional (including cell) differentiation.     

Neuronal avalanches of a self-organized neural network with active-neuron-dominant structure

Neuronal avalanche is a spontaneous neuronal activity which obeys a power-law distribution of population event sizes with an exponent of -3/2. It has been observed in the superficial layers of cortex both in vivo and in vitro. In this paper, we analyze the information transmission of a novel self-organized neural network with active-neuron-dominant structure. Neuronal avalanches can be observed in this network with appropriate input intensity. We find that the process of network learning via spike-timing dependent plasticity dramatically increases the complexity of network structure, which is finally self-organized to be active-neuron-dominant connectivity. Both the entropy of activity patterns and the complexity of their resulting post-synaptic inputs are maximized when the network dynamics are propagated as neuronal avalanches. This emergent topology is beneficial for information transmission with high efficiency and also could be responsible for the large information capacity of this network compared with alternative archetypal networks with different neural connectivity.     

Coherent optical neural networks and the generalization characteristic

Coherent optical neural-network systems are presented by which we are to realize (1) control of optical neural-network behavior by carrier frequency modulation and (2) frequency-domain multiplexing as a new degree of neural parallelism. In the coherent optical neural network, lightwave carries amplitude, phase, and frequency signals which are processed through optical neural connections consisting of transparency, delay, and optical nonlinear neurons. The neural learning process is realized by adjusting the delay time and the transpar-ency of the optical connections by regarding the carrier frequency as a learning parameter. A simulation experiment demonstrates that the proposed system learns desired output signals depending on the optical carrier frequencies, which leads to the above advantages. It is also found that the generalization characteristics depends on built-in input delays. The generalization quality is discussed from the viewpoint of function approximation or synthesis using sinusoidal basis functions.     

The influence of connectivity on the firing rate in a neuronal network with electrical and chemical synapses

We study the firing rate properties of a cellular automaton model for a neuronal network with chemical synapses. We propose a simple mechanism in which the nonlocal connections are included, through electrical and chemical synapses. In the latter case, we introduce a time delay which produces self-sustained activity. Nonlocal connections, or shortcuts, are randomly introduced according to a specified connection probability. There is a range of connection probabilities for which neuron firing occurs, as well as a critical probability for which the firing ceases in the absence of time delay. The critical probability for nonlocal shortcuts depends on the network size according to a power-law. We also compute the firing rate amplification factor by varying both the connection probability and the time delay for different network sizes.     

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.     

Nonlinear transfer function encodes synchronization in a neural network from the mammalian brain

Synchronization is one of the mechanisms by which the brain encodes information. The observed synchronization of neuronal activity has, however, several levels of fluctuations, which presumably regulate local features of specific areas. This means that biological neural networks should have an intrinsic mechanism able to synchronize the neuronal activity but also to preserve the firing capability of individual cells. Here, we investigate the input-output relationship of a biological neural network from developing mammalian brain, i.e., the hippocampus. We show that the probability of occurrence of synchronous output activity (which consists in stereotyped population bursts recorded throughout the hippocampus) is encoded by a sigmoidal transfer function of the input frequency. Under this scope, low-frequency inputs will not produce any coherent output while high-frequency inputs will determine a synchronous pattern of output activity (population bursts). We analyze the effect of the network size (N) on the parameters of the transfer function (threshold and slope). We found that sigmoidal functions realistically simulate the synchronous output activity of hippocampal neural networks. This outcome is particularly important in the application of results from neural network models to neurobiology.     

The influence of hubs in the structure of a neuronal network during an epileptic seizure

In this work, we propose changes in the structure of a neuronal network with the intention to provoke strong synchronization to simulate episodes of epileptic seizure. Starting with a network of Izhikevich neurons we slowly increase the number of connections in selected nodes in a controlled way, to produce (or not) hubs. We study how these structures alter the synchronization on the spike firings interval, on individual neurons as well as on mean values, as a function of the concentration of connections for random and non-random (hubs) distribution. We also analyze how the post-ictal signal varies for the different distributions. We conclude that a network with hubs is more appropriate to represent an epileptic state.     

The growth of cognition: Free energy minimization and the embryogenesis of cortical computation

The assumption that during cortical embryogenesis neurons and synaptic connections are selected to form an ensemble maximising synchronous oscillation explains mesoscopic cortical development, and a mechanism for cortical information processing is implied by consistency with the Free Energy Principle and Dynamic Logic. A heteroclinic network emerges, with stable and unstable fixed points of oscillation corresponding to activity in symmetrically connected, versus asymmetrically connected, sets of neurons. Simulations of growth explain a wide range of anatomical observations for columnar and non-columnar cortex, superficial patch connections, and the organization and dynamic interactions of neurone response properties. An antenatal scaffold is created, upon which postnatal learning can establish continuously ordered neuronal representations, permitting matching of co-synchronous fields in multiple cortical areas to solve optimization problems as in Dynamic Logic. Fast synaptic competition partitions equilibria, minimizing “the curse of dimensionality”, while perturbations between imperfectly partitioned synchronous fields, under internal reinforcement, enable the cortex to become adaptively self-directed. As learning progresses variational free energy is minimized and entropy bounded.     

Plasticity-induced characteristic changes of pattern dynamics and the related phase transitions in small-world neuronal networks

Phase transitions widely exist in nature and occur when some control parameters are changed. In neural systems, their macroscopic states are represented by the activity states of neuron populations, and phase transitions between different activity states are closely related to corresponding functions in the brain. In particular, phase transitions to some rhythmic synchronous firing states play significant roles on diverse brain functions and disfunctions, such as encoding rhythmical external stimuli, epileptic seizure, etc. However, in previous studies, phase transitions in neuronal networks are almost driven by network parameters （e.g., external stimuli）, and there has been no investigation about the transitions between typical activity states of neuronal networks in a self-organized way by applying plastic connection weights. In this paper, we discuss phase transitions in electrically coupled and lattice-based small-world neuronal networks （LBSW networks） under spike-timing-dependent plasticity （STDP）. By applying STDP on all electrical synapses, various known and novel phase transitions could emerge in LBSW networks, particularly, the phenomenon of self-organized phase transitions （SOPTs）： repeated transitions between synchronous and asynchronous firing states. We further explore the mechanics generating SOPTs on the basis of synaptic weight dynamics.     

Analysis of spontaneous activity in neuronal cultures through recurrence plots: impact of varying connectivity

We analyzed spontaneous activity of cortical neuronal networks in vitro using recurrence plots (RPs). Our data encompasses fluorescence traces of average network activity from two experimental explorations, namely the development of connections during the maturation of the network and the gradual weakening of connections through chemical action. The dynamical richness of the networks in these connectivity-evolving scenarios was examined through recurrence quantification analysis. Measures such as determinism and laminarity were used to portray the degree of uniformity and periodicity of the spontaneous activity patterns. The analysis shows that RPs are a powerful tool to visualize and interpret neuronal networks dynamics, and pinpoint its hallmarks.     

Multilayer adaptive networks in neuronal processing

The connectome is a wiring diagram mapping all the neural connections in the brain. At the cellular level, it provides a map of the neurons and synapses within a part or all of the brain of an organism. In recent years, significant advances have been made in the study of the connectome via network science and graph theory. This analysis is fundamental to understand neurotransmission (fast synaptic transmission) networks. However, neurons use other forms of communication as neuromodulation that, instead of conveying excitation or inhibition, change neuronal and synaptic properties. This additional neuromodulatory layers condition and reconfigure the connectome. In this paper, we propose that multilayer adaptive networks, in which different synaptic and neurochemical layers interact, are the appropriate framework to explain neuronal processing. Then, we describe a simplified multilayer adaptive network model that accounts for these extra-layers of interaction and analyse the emergence of interesting computational capabilities.     

Dynamical encoding by networks of competing neuron groups: winnerless competition

Following studies of olfactory processing in insects and fish, we investigate neural networks whose dynamics in phase space is represented by orbits near the heteroclinic connections between saddle regions (fixed points or limit cycles). These networks encode input information as trajectories along the heteroclinic connections. If there are N neurons in the network, the capacity is approximately e(N-1)!, i.e., much larger than that of most traditional network structures. We show that a small winnerless competition network composed of FitzHugh-Nagumo spiking neurons efficiently transforms input information into a spatiotemporal output.     

Synchronizability of stochastic network ensembles in a model of interacting dynamical units

Synchronization is an important phenomenon which occurs in the dynamics of complex systems. Synchronized states emerge both from an adjustment of the system parameters and from an application of a proper external stimulus. In the present paper we study synchronized activity in a neural network model whose dynamics is driven by an external activation. In this context we are interested in its synchronizability, i.e. the existence of inputs causing the model system to synchronize. Furthermore, we investigate global synchronizability properties of stochastic network structure ensembles (instead of single realizations of a network architecture). We study the small world network, a model of preferential linking structure, and the classical Erd?s-Renyi random graph as particular examples of network topologies. Their synchronizability properties are investigated by analytical arguments and numerical simulations. Our analysis shows the emergence of synchronizable states of network ensembles for a wide range of the parameter values. In addition we observe and study the transition behaviour from synchronizability to nonsynchronizability.     

Communication on the structure of biological networks

Networks are widely used to represent interaction pattern among the components in complex systems. Structures of real networks from different domains may vary quite significantly. As there is an interplay between network architecture and dynamics, structure plays an important role in communication and spreading of information in a network. Here we investigate the underlying undirected topology of different biological networks which support faster spreading of information and are better in communication. We analyse the good expansion property by using the spectral gap and communicability between nodes. Different epidemic models are also used to study the transmission of information in terms of spreading of disease through individuals (nodes) in those networks. Moreover, we explore the structural conformation and properties which may be responsible for better communication. Among all biological networks studied here, the undirected structure of neuronal networks not only possesses the small-world property but the same is also expressed remarkably to a higher degree compared to any randomly generated network which possesses the same degree sequence. A relatively high percentage of nodes, in neuronal networks, form a higher core in their structure. Our study shows that the underlying undirected topology in neuronal networks, in a significant way, is qualitatively different from the same in other biological networks and that they may have evolved in such a way that they inherit a (undirected) structure which is excellent and robust in communication.     

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.