Synaptic, dendritic and single-cell kinetics generate significant time delays that shape the dynamics of large networks of spiking neurons. Previous work has shown that such effective delays can be taken into account with a rate model through the addition of an explicit, fixed delay [Roxin et al. PRL 238103 (2005)]. Here we extend this work to account for arbitrary symmetric patterns of synaptic connectivity and generic nonlinear transfer functions. Specifically, we conduct a weakly nonlinear analysis of the dynamical states arising via primary instabilities of the asynchronous state. In this way we determine analytically how the nature and stability of these states depend on the choice of transfer function and connectivity. We arrive at two general observations of physiological relevance that could not be explained in previous works. These are: 1 - Fast oscillations are always supercritical for realistic transfer functions. 2 - Traveling waves are preferred over standing waves given plausible patterns of local connectivity. We finally demonstrate that these results show a good agreement with those obtained performing numerical simulations of a network of Hodgkin-Huxley neurons.
We show that the unavoidable increase in neuronal response latency to ongoing stimulation serves as a nonuniform gradual stretching of neuronal circuit delay loops and emerges as an essential mechanism in the formation of various types of neuronal timers. Synchronization emerges as a transient phenomenon without predefined precise matched synaptic delays. These findings are described in an experimental procedure where conditioned stimulations were enforced on a circuit of neurons embedded within a large-scale network of cortical cells in-vitro, and are corroborated by neuronal simulations. The…
Propagation of oscillatory signals through the cortex and coherence is shaped by the connectivity structure of neuronal circuits. This study systematically investigates the network and stimulus properties that shape network responses. The results show how input to a cortical column model of the primary visual cortex excites dynamical modes determined by the laminar pattern. Stimulating the inhibitory neurons in the upper layer reproduces experimentally observed resonances at gamma frequency whose origin can be traced back to two anatomical sub-circuits. We develop this result systematically: I…
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 study the time delay in the synaptic conductance for suppression of spike synchronisation in a random network of Hodgkin Huxley neurons coupled by means of chemical synapses. In the first part, we examine in detail how the time delay acts over the network during the synchronised and desynchronised neuronal activities. We observe a relation between the neuronal dynamics and the syaptic conductance distributions. We find parameter values in which the time delay has high effectiveness in promoting the suppression of spike synchronisation. In the second part, we analyse how the delayed neuronal…
Oscillatory activities are widely observed in specific frequency bands of recorded field potentials in different brain regions, and play critical roles in processing neural information. Understanding the structure of these oscillatory activities is essential for understanding the brain function. So far many details remain elusive about their rhythmic structures and how these oscillations are generated. We show that many oscillatory activities in spontaneous cortical-subcortical networks, such as delta, spindle, gamma, high-gamma and sharp wave ripple bands in different brain regions, are genui…
We present two novel methods for performing logic operations. Our methods are based on using the time dimension for programming and data representation. The first method is based on varying the sampling moment in time of a neuronal action potential, and the second method is based on a neural delay system, where the generation of the action potential is delayed by specific time lengths, to be sampled at a fixed moment in time. Both methods are supported by explicit examples.
This paper investigates the dependence of synchronization transitions of bursting oscillations on the information transmission delay over scale-free neuronal networks with attractive and repulsive coupling. It is shown that for both types of coupling, the delay always plays a subtle role in either promoting or impairing synchronization. In particular, depending on the inherent oscillation period of individual neurons, regions of irregular and regular propagating excitatory fronts appear intermittently as the delay increases. These delay-induced synchronization transitions are manifested as wel…
We study the effects of periodic subthreshold pacemaker activity and time-delayed coupling on stochastic resonance over scale-free neuronal networks. As the two extreme options, we introduce the pacemaker respectively to the neuron with the highest degree and to one of the neurons with the lowest degree within the network, but we also consider the case when all neurons are exposed to the periodic forcing. In the absence of delay, we show that an intermediate intensity of noise is able to optimally assist the pacemaker in imposing its rhythm on the whole ensemble, irrespective to its placing, t…
Realistic networks display heterogeneous transmission delays. We analyze here the limits of large stochastic multi-populations networks with stochastic coupling and random interconnection delays. We show that depending on the nature of the delays distributions, a quenched or averaged propagation of chaos takes place in these networks, and that the network equations converge towards a delayed McKean-Vlasov equation with distributed delays. Our approach is mostly fitted to neuroscience applications. We instantiate in particular a classical neuronal model, the Wilson and Cowan system, and show th…
We investigate front propagation and synchronization transitions in dependence on the information transmission delay and coupling strength over scale-free neuronal networks with different average degrees and scaling exponents. As the underlying model of neuronal dynamics, we use the efficient Rulkov map with additive noise. We show that increasing the coupling strength enhances synchronization monotonously, whereas delay plays a more subtle role. In particular, we found that depending on the inherent oscillation frequency of individual neurons, regions of irregular and regular propagating exci…
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…
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…
A recurrent loop consisting of a single neuron is considered which is influenced by a chemical excitatory delayed synaptic feedback. We show the response of the system is dependent to the duration of the activity of the synapse which is determined by the deactivation time constant of the synapse. We show that loops with slow synapses, those which the effect of the synaptic activation remains for time constants comparable to the period of firing, show more predictable results where the effect of the fast synapses is tightly dependent on the loop delay time. The results are compared to those of …
In this paper, we propose a dynamic delayed feedback control approach for desynchronization of chaotic-bursting synchronous activities in an ensemble of globally coupled neuronal oscillators. We demonstrate that the difference signal between an ensemble's mean field and its time delayed state, filtered and fed back to the ensemble, can suppress the self-synchronization in the ensemble. These individual units are decoupled and stabilized at the desired desynchronized states while the stimulation signal reduces to the noise level. The effectiveness of the method is illustrated by examples of two…
Here we investigate the synchronization of networks of FitzHugh-Nagumo neurons coupled in scale-free, small-world and random topologies, in the presence of distributed time delays in the coupling of neurons. We explore how the synchronization transition is affected when the time delays in the interactions between pairs of interacting neurons are non-uniform. We find that the presence of distributed time-delays does not change the behavior of the synchronization transition significantly, vis-a-vis networks with constant time-delay, where the value of the constant time-delay is the mean of the d…
First spike latency following stimulus onset is of significant physiological relevance. Neurons transmit information about their inputs by transforming them into spike trains, and the timing of these spike trains is in turn crucial for effectively encoding that information. Random processes and uncertainty that underly neuronal dynamics have been shown to prolong the time towards the first response in a phenomenon dubbed noise-delayed decay. Here we study whether Hodgkin-Huxley neurons with a tunable intensity of intrinsic noise might have shorter response times to external stimuli just above …
We discuss synchronization in networks of neuronal oscillators which are interconnected via diffusive coupling, i.e. linearly coupled via gap junctions. In particular, we present sufficient conditions for synchronization in these networks using the theory of semi-passive and passive systems. We show that the conductance-based neuronal models of Hodgkin-Huxley, Morris-Lecar, and the popular reduced models of FitzHugh-Nagumo and Hindmarsh-Rose all satisfy a semi-passivity property, i.e. that is the state trajectories of such a model remain oscillatory but bounded provided that the supplied (elec…
Networks of the brain are composed of a very large number of neurons connected through a random graph and interacting after random delays that both depend on the anatomical distance between cells. In order to comprehend the role of these random architectures on the dynamics of such networks, we analyze the mesoscopic and macroscopic limits of networks with random correlated connectivity weights and delays. We address both averaged and quenched limits, and show propagation of chaos and convergence to a complex integral McKean-Vlasov equations with distributed delays. We then instantiate a compl…
We consider a simple Markovian class of the stochastic Wilson-Cowan type models of neuronal network dynamics, which incorporates stochastic delay caused by the existence of a refractory period of neurons. From the point of view of the dynamics of the individual elements, we are dealing with a network of non-Markovian stochastic two-state oscillators with memory which are coupled globally in a mean-field fashion. This interrelation of a higher-dimensional Markovian and lower-dimensional non-Markovian dynamics is discussed in its relevance to the general problem of the network dynamics of comple…
Networks of fast-spiking interneurons are crucial for the generation of neural oscillations in the brain. Here we study the synchronous behavior of interneuronal networks that are coupled by delayed inhibitory and fast electrical synapses. We find that both coupling modes play a crucial role by the synchronization of the network. In addition, delayed inhibitory synapses affect the emerging oscillatory patterns. By increasing the inhibitory synaptic delay, we observe a transition from regular to mixed oscillatory patterns at a critical value. We also examine how the unreliability of inhibitory …
Analytical expressions for scaling of brain wave spectra derived from the general nonlinear wave Hamiltonian form show excellent agreement with experimental "neuronal avalanche" data. The theory of the weakly evanescent nonlinear brain wave dynamics reveals the underlying collective processes hidden behind the phenomenological statistical description of the neuronal avalanches and connects together the whole range of brain activity states, from oscillatory wave-like modes, to neuronal avalanches, to incoherent spiking, showing that the neuronal avalanches are just the manifestation of the diff…
Oscillatory dynamics are ubiquitous in biological networks. Possible sources of oscillations are well understood in low-dimensional systems, but have not been fully explored in high-dimensional networks. Here we study large networks consisting of randomly coupled rate units. We identify a novel type of bifurcation in which a continuous part of the eigenvalue spectrum of the linear stability matrix crosses the instability line at non-zero-frequency. This bifurcation occurs when the interactions are delayed and partially anti-symmetric, and leads to a heterogeneous oscillatory state in which osc…
The instantaneous state of a neural network consists of both the degree of excitation of each neuron, the network is composed of, and positions of impulses in communication lines between neurons. In neurophysiological experiments, the neuronal firing moments are registered, but not the state of communication lines. But future spiking moments depend essentially on the past positions of impulses in the lines. This suggests, that the sequence of intervals between firing moments (interspike intervals, ISIs) in the network could be non-Markovian. In this paper, we address this question for a simple…
Synchronized bursts (SBs) with complex structures are common in neuronal cultures. Although the origin of SBs is still unclear, they have been studied for their information processing capabilities. Here, we investigate the properties of these SBs in a culture on multi-electrode array system. We find that structures of these SBs are related to the different developmental stages of the cultures. A model based on short term synaptic plasticity, recurrent connections and astrocytic recycling of neurotransmitters has been developed successfully to understand these structures. A phase diagram obtain…
We present a use of modern data-based machine learning approaches to suppress self-sustained collective oscillations typically signaled by ensembles of degenerative neurons in the brain. The proposed hybrid model relies on two major components: an environment of oscillators and a policy-based reinforcement learning block. We report a model-agnostic synchrony control based on proximal policy optimization and two artificial neural networks in an Actor-Critic configuration. A class of physically meaningful reward functions enabling the suppression of collective oscillatory mode is proposed. The s…
Neurons in many brain areas can develop pronounced depolarized state of membrane potential (up state) in addition to the normal hyperpolarized down state near the resting potential. The influence of the up state on signal encoding, however, is not well investigated. Here we construct a one-dimensional bistable neuron model and calculate the linear response to noisy oscillatory inputs analytically. We find that with the appearance of an up state, the transmission function is enhanced by the emergence of a local maximum at some optimal frequency and the phase lag relative to the input signal is …
Synaptic delays modulate population phase and amplitude responses in oscillatory excitatory-inhibitory networks
Institute for Advanced Studies in Basic Sciences (IASBS)（基础科学高等研究院）；Institute for Research in Fundamental Sciences (IPM)（基本科学研究院）；Pasargad Institute for Advanced Innovative Solutions (PIAIS)（帕萨尔加德先进创新解决方案研究所）；The Zapata-Briceño Institute of Neuroscience（萨帕塔-布里斯尼奥神经科学研究所）
Brain plasticity refers to brain's ability to change neuronal connections, as a result of environmental stimuli, new experiences, or damage. In this work, we study the effects of the synaptic delay on both the coupling strengths and synchronisation in a neuronal network with synaptic plasticity. We build a network of Hodgkin-Huxley neurons, where the plasticity is given by the Hebbian rules. We verify that without time delay the excitatory synapses became stronger from the high frequency to low frequency neurons and the inhibitory synapses increases in the opposite way, when the delay is incre…
We numerically investigate the influence of intrinsic channel noise on the dynamical response of delay-coupling in neuronal systems. The stochastic dynamics of the spiking is modeled within a stochastic modification of the standard Hodgkin-Huxley model wherein the delay-coupling accounts for the finite propagation time of an action potential along the neuronal axon. We quantify this delay-coupling of the Pyragas-type in terms of the difference between corresponding presynaptic and postsynaptic membrane potentials. For an elementary neuronal network consisting of two coupled neurons we detect c…
We propose a new experimentally corroborated paradigm in which the functionality of the brain's logic-gates depends on the history of their activity, e.g. an OR-gate that turns into a XOR-gate over time. Our results are based on an experimental procedure where conditioned stimulations were enforced on circuits of neurons embedded within a large-scale network of cortical cells in-vitro. The underlying biological mechanism is the unavoidable increase of neuronal response latency to ongoing stimulations, which imposes a non-uniform gradual stretching of network delays.
A model of the columnar functional organization of neocortical association areas is studied. The neuronal network is composed of many Hebbian autoassociators, or modules, each of which interacts with a relatively small number of the others. Every module encodes and stores a number of elementary percepts, or features. Memory items, or patterns, are peculiar combinations of features sparsely distributed over the multi-modular network. Any feature stored in any module can be involved in several of the stored patterns; feature-sharing is in fact source of local ambiguities and, consequently, a pot…
We have studied effects of perturbations on the cat cerebral cortex. According to the literature, this cortex structure can be described by a clustered network. This way, we construct a clustered network with the same number of areas as in the cat matrix, where each area is described as a sub-network with small-world property. We focus on the suppression of neuronal phase synchronisation considering different kinds of perturbations. Among the various controlling interventions, we choose three methods: delayed feedback control, external time-periodic driving, and activation of selected neurons.…
The oculomotor delayed-response (ODR) task is a common experimental paradigm of working memory (WM) study, in which a monkey must fixate its gaze on the center of a screen and, following a brief cue that flashes on the screen, keep fixating for several more seconds before shifting its gaze to the location where the cue flashed. Consequently, in the delay period between the cue and the response the monkey must maintain a memory of cue location. Electrophysiological recordings from the prefrontal area of the cortex (PFC) revealed neurons that display selective persistent activity: their firing r…
Synaptic connections in neuronal circuits are modulated by pre- and post-synaptic spiking activity. Heuristic models of this process of synaptic plasticity can provide excellent fits to results from in-vitro experiments in which pre- and post-synaptic spiking is varied in a controlled fashion. However, the plasticity rules inferred from fitting such data are inevitably unstable, in that given constant pre- and post-synaptic activity the synapse will either fully potentiate or depress. This instability can be held in check by adding additional mechanisms, such as homeostasis. Here we consider a…
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 ob…
In 1943 McCulloch and Pitts suggested that the brain is composed of reliable logic-gates similar to the logic at the core of today's computers. This framework had a limited impact on neuroscience, since neurons exhibit far richer dynamics. Here we propose a new experimentally corroborated paradigm in which the truth tables of the brain's logic-gates are time dependent, i.e. dynamic logicgates (DLGs). The truth tables of the DLGs depend on the history of their activity and the stimulation frequencies of their input neurons. Our experimental results are based on a procedure where conditioned sti…
Delay coordinates synchronization and induces abrupt transition in excitable networks
Institute of Science and Technology, Universidade Federal de São Paulo (UNIFESP)（圣保罗联邦大学科学与技术学院）；Cellular Computations and Learning, Max Planck Institute for Neurobiology of Behavior（马克斯·普朗克行为神经生物学研究所细胞计算与学习部门）；Department of Mathematics, Western University（韦仕敦大学数学系）；Fields Lab for Network Computation, Fields Institute（菲尔兹研究所网络计算实验室）；Department of Neuroscience, University of Lethbridge（莱斯布里奇大学生物神经科学系）
Regime Mapping of Oscillatory States in Balanced Spiking Networks with Multiple Time Scales
Graduate Institute of Networking and Multimedia, National Taiwan University（台湾大学网际多媒体研究所）；Graduate Institute of Vehicle Engineering, National Changhua University of Education（国立彰化师范大学车辆工程研究所）
The conventional impedance profile of a neuron can identify the presence of resonance and other properties of the neuronal response to oscillatory inputs, such as nonlinear response amplifications, but it cannot distinguish other nonlinear properties such as asymmetries in the shape of the voltage response envelope. Experimental observations have shown that the response of neurons to oscillatory inputs preferentially enhances either the upper or lower part of the voltage envelope in different frequency bands. These asymmetric voltage responses arise in a neuron model when it is submitted to hi…
Collective chaos is shown to emerge, via a period-doubling cascade, from quasiperiodic partial synchronization in a population of identical inhibitory neurons with delayed global coupling. This system is thoroughly investigated by means of an exact model of the macroscopic dynamics, valid in the thermodynamic limit. The collective chaotic state is reproduced numerically with a finite population, and persists in the presence of weak heterogeneities. Finally, the relationship of the model's dynamics with fast neuronal oscillations is discussed.
Activity in coupled systems is often oscillatory, for example, the firing pattern of neuronal populations. Whereas these oscillations have been studied predominantly in local circuits, here we show how the topology of large-scale networks, leading to large feedback loops, influences oscillations in the resting state. We find that the hierarchical modular organization of neuronal networks supports distinct spectral patterns of neural rhythms similar to those observed experimentally in different species such as rat and human. For individual neurons, multiple peak frequencies with non-integer rat…
INTENSE: Detecting and disentangling neuronal selectivity in calcium imaging data
Institute for Advanced Brain Studies of the Moscow State University（莫斯科国立大学高级脑科学研究所）；Center for Interdisciplinary Research in Biology, Collège de France（法兰西公学院跨学科生物学研究中心）；Semenov Institute of Chemical Physics, Russian Academy of Sciences（俄罗斯科学院谢苗诺夫化学物理研究所）
Persistent activity is postulated to drive neural network plasticity and learning. To investigate its underlying cellular mechanisms, we developed a biophysically tractable model that explains the emergence, sustenance, and eventual termination of short-term persistent activity. Using the model, we reproduced the features of reverberating activity that were observed in small (50-100 cells) networks of cultured hippocampal neurons, such as the appearance of polysynaptic current clusters, the typical inter-cluster intervals, the typical duration of reverberation, and the response to changes in e…
When inhibitory neurons constitute about 40% of neurons they could have an important antinociceptive role, as they would easily regulate the level of activity of other neurons. We consider a simple network of cortical spiking neurons with axonal conduction delays and spike timing dependent plasticity, representative of a cortical column or hypercolumn with large proportion of inhibitory neurons. Each neuron fires following a Hodgkin-Huxley like dynamics and it is interconnected randomly to other neurons. The network dynamics is investigated estimating Bandt and Pompe probability distribution f…
A class of spiking neuronal models with threshold 2 is considered. It is defined by a set of conditions typical for basic threshold-type models, such as the leaky integrate-and-fire (LIF) or the binding neuron model and also for some artificial neurons. A neuron is stimulated with a Poisson stream of excitatory impulses. Each output impulse is conveyed through the feedback line to the neuron input after finite delay $Δ$. This impulse is identical to those delivered from the input stream. We have obtained a general relation allowing calculating exactly the probability density function (PDF) $p(…
