This review gives a short historical account of the excitable maps approach for modeling neurons and neuronal networks. Some early models, due to Pasemann (1993), Chialvo (1995) and Kinouchi and Tragtenberg (1996), are compared with more recent proposals by Rulkov (2002) and Izhikevich (2003). We also review map-based schemes for electrical and chemical synapses and some recent findings as critical avalanches in map-based neural networks. We conclude with suggestions for further work in this area like more efficient maps, compartmental modeling and close dynamical comparison with conductance-based models.
We study the dynamical states that emerge in a small-world network of recurrently coupled excitable neurons through both numerical and analytical methods. These dynamics depend in large part on the fraction of long-range connections or `short-cuts' and the delay in the neuronal interactions. Persistent activity arises for a small fraction of `short-cuts', while a transition to failure occurs at a critical value of the `short-cut' density. The persistent activity consists of multi-stable periodic attractors, the number of which is at least on the order of the number of neurons in the network. F…
We demonstrate the existence of noise-induced periodicity (coherence resonance) in both a discrete-time model and a continuous-time model of an excitable neuron. In particular, we show that the effects of noise added to the fast and slow dynamics of the models are dramatically different. A Fokker-Planck analysis gives a quantitative explanation of the effects.
We study the dynamics of excitable integrate-and-fire neurons in a small-world network. At low densities $p$ of directed random connections, a localized transient stimulus results in either self-sustained persistent activity or in a brief transient followed by failure. Averages over the quenched ensemble reveal that the probability of failure changes from 0 to 1 over a narrow range in $p$; this failure transition can be described analytically through an extension of an existing mean-field result. Exceedingly long transients emerge at higher densities $p$; their activity patterns are disordered…
We present a new interpretation for encoding information of the period of input signals into spike-trains in individual sensory neuronal systems. The spike-train could be described as the waveform sample of the input signal which locks sample points to wave crests with randomness. Based on simulations of the Hodgkin-Huxley (HH) neuron responding to periodic inputs, we demonstrate that the random sampling is a proper encoding method in medium frequency region since power spectra of the reconstructed spike-trains are identical to that of neural signals.
The emergence and stability of splay states is studied in fully coupled finite networks of N excitable quadratic integrate-and-fire neurons, connected via synapses modeled as pulses of finite amplitude and duration. For such synapses, by introducing two distinct types of synaptic events (pulse emission and termination), we were able to write down an exact event-driven map for the system and to evaluate the splay state solutions. For M overlapping post synaptic potentials the linear stability analysis of the splay state should take in account, besides the actual values of the membrane potential…
An electronic circuit device, inspired on the FitzHugh-Nagumo model of neuronal excitability, was constructed and shown to operate with characteristics compatible with those of biological sensory neurons. The nonlinear dynamical model of the electronics quantitatively reproduces the experimental observations on the circuit, including the Hopf bifurcation at the onset of tonic spiking. Moreover, we have implemented an analog noise generator as a source to study the variability of the spike trains. When the circuit is in the excitable regime, coherence resonance is observed. At sufficiently low …
In this work, we study the dynamic range of a neuronal network of excitable neurons with excitatory and inhibitory synapses. We obtain an analytical expression for the critical point as a function of the excitatory and inhibitory synaptic intensities. We also determine an analytical expression that gives the critical point value in which the maximal dynamic range occurs. Depending on the mean connection degree and coupl\-ing weights, the critical points can exhibit ceasing or ceaseless dynamics. However, the dynamic range is equal in both cases. We observe that the external stimulus mask some …
According to the conventional approach to model neural ensembles the extracellular environment has fixed ionic concentrations. However, in many cases the extracellular concentration of potassium ions can not be regarded as constant. That represents specific chemical pathway for neurons to interact and can influence strongly the behavior of a single neuron as well as large ensembles. The released chemical agent follows a diffusive dynamics in the external medium, that lowers the threshold of individual excitable units. We address this problem by studying simplified excitable units given by a mo…
The study of balanced networks of excitatory and inhibitory neurons has led to several open questions. On the one hand it is yet unclear whether the asynchronous state observed in the brain is autonomously generated, or if it results from the interplay between external drivings and internal dynamics. It is also not known, which kind of network variabilities will lead to irregular spiking and which to synchronous firing states. Here we show how isolated networks of purely excitatory neurons generically show asynchronous firing whenever a minimal level of structural variability is present togeth…
Spontaneous formation of clusters of synchronized spiking in a structureless ensemble of equal stochastically perturbed excitable neurons with delayed coupling is demonstrated for the first time. The effect is a consequence of a subtle interplay between interaction delays, noise and the excitable character of a single neuron. Dependence of the cluster properties on the time-lag, noise intensity and the synaptic strength is investigated.
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）
This article communicates results on regular depolarization cascades in periodically-kicked feedforward chains of excitable two-dimensional FitzHugh-Nagumo systems driven by sufficiently strong excitatory forcing at the front node. The study documents a parameter exploration by way of changes to the forcing period, upon which the dynamics undergoes a transition from simple depolarization to more complex behavior, including the emergence of mixed-mode oscillations. Both rigorous studies and careful numerical observations are presented. In particular, we provide rigorous proofs for existence and…
The scaling of transient times to zero-lag synchronization in networks composed of excitable units is shown to be governed by three features of the graph representing the network: the longest path between pairs of neurons (diameter), the largest loop (circumference) and the loop with the maximal average out degree. The upper bound of transient times can vary between O(1) and O(N2), where N is the size of the network, and its scaling can be predicted in many scenarios from finite time accumulated information of the transient. Results challenge the assumption that functionality of neural network…
A recent study on the effect of colored driving noise on the escape from a metastable state derives an analytic expression of the transfer function of the leaky integrate-and-fire neuron model subject to colored noise. Here we present an alternative derivation of the results, taking into account time-dependent boundary conditions explicitly. This systematic approach may facilitate future extensions beyond first order perturbation theory. The analogy of the quantum harmonic oscillator to the LIF neuron model subject to white noise enables a derivation of the well known transfer function simpler…
We study a simple map as a minimal model of excitable cells. The map has two fast variables which mimic the behavior of class I neurons, undergoing a sub-critical Hopf bifurcation. Adding a third slow variable allows the system to present bursts and other interesting biological behaviors. Bifurcation lines which locate the excitability region are obtained for different planes in parameter space.
In sensory neurons the presence of noise can facilitate the detection of weak information-carrying signals, which are encoded and transmitted via correlated sequences of spikes. Here we investigate relative temporal order in spike sequences induced by a subthreshold periodic input, in the presence of white Gaussian noise. To simulate the spikes, we use the FitzHugh-Nagumo model, and to investigate the output sequence of inter-spike intervals (ISIs), we use the symbolic method of ordinal analysis. We find different types of relative temporal order, in the form of preferred ordinal patterns whic…
As few real systems comprise indistinguishable units, diversity is a hallmark of nature. Diversity among interacting units shapes properties of collective behavior such as synchronization and information transmission. However, the benefits of diversity on information processing at the edge of a phase transition, ordinarily assumed to emerge from identical elements, remain largely unexplored. Analyzing a general model of excitable systems with heterogeneous excitability, we find that diversity can greatly enhance optimal performance (by two orders of magnitude) when distinguishing incoming inpu…
We show that \emph{stochastic bursting} is observed in a ring of unidirectional delay-coupled noisy excitable systems, thanks to the combinational action of time-delayed coupling and noise. Under the approximation of timescale separation, i.e., when the time delays in each connection are much larger than the characteristic duration of the spikes, the observed rather coherent spike pattern can be described by idealized coupled point processes with a leader-follower relationship. We derive analytically the statistics of the spikes in each unit, pairwise correlations between any two units, and th…
In the light of recent experimental findings that gap junctions are essential for low level intensity detection in the sensory periphery, the Greenberg-Hastings cellular automaton is employed to model the response of a two-dimensional sensory network to external stimuli. We show that excitable elements (sensory neurons) that have a small dynamical range are shown to give rise to a collective large dynamical range. Therefore the network transfer (gain) function (which is Hill or Stevens law-like) is an emergent property generated from a pool of small dynamical range cells, providing a basis for…
When a simple excitable system is continuously stimulated by a Poissonian external source, the response function (mean activity versus stimulus rate) generally shows a linear saturating shape. This is experimentally verified in some classes of sensory neurons, which accordingly present a small dynamic range (defined as the interval of stimulus intensity which can be appropriately coded by the mean activity of the excitable element), usually about one or two decades only. The brain, on the other hand, can handle a significantly broader range of stimulus intensity, and a collective phenomenon in…
We study the statistical physics of a surprising phenomenon arising in large networks of excitable elements in response to noise: while at low noise, solutions remain in the vicinity of the resting state and large-noise solutions show asynchronous activity, the network displays orderly, perfectly synchronized periodic responses at intermediate level of noise. We show that this phenomenon is fundamentally stochastic and collective in nature. Indeed, for noise and coupling within specific ranges, an asymmetry in the transition rates between a resting and an excited regime progressively builds up…
The dynamic range measures the capacity of a system to discriminate the intensity of an external stimulus. Such an ability is fundamental for living beings to survive: to leverage resources and to avoid danger. Consequently, the larger is the dynamic range, the greater is the probability of survival. We investigate how the integration of different input signals affects the dynamic range, and in general the collective behavior of a network of excitable units. By means of numerical simulations and a mean-field approach, we explore the nonequilibrium phase transition in the presence of integratio…
We study the stochastic dynamics of strongly-coupled excitable elements on a tree network. The peripheral nodes receive independent random inputs which may induce large spiking events propagating through the branches of the tree and leading to global coherent oscillations in the network. This scenario may be relevant to action potential generation in certain sensory neurons, which possess myelinated distal dendritic tree-like arbors with excitable nodes of Ranvier at peripheral and branching nodes and exhibit noisy periodic sequences of action potentials. We focus on the spiking statistics of …
Influence of mesoscopic channel noise on excitable dynamics of living cells became a hot subject within the last decade, and the traditional biophysical models of neuronal dynamics such as Hodgkin-Huxley model have been generalized to incorporate such effects. There still exists but a controversy on how to do it in a proper and computationally efficient way. Here we introduce an improved Langevin description of stochastic Hodgkin-Huxley dynamics with natural boundary conditions for gating variables. It consistently describes the channel noise variance in a good agreement with discrete state mo…
We study the emergence and coherence of stochastic oscillations in star networks of excitable elements in which peripheral nodes receive independent random inputs. A biophysical model of a distal branch of sensory neuron in which peripheral nodes of Ranvier are coupled to a central node by myelinated cable segments is used along with a generic model of networked stochastic active rotators. We show that coherent oscillations can emerge due to stochastic synchronization of peripheral nodes and that the degree of coherence can be maximized by tuning the coupling strength and the size of the netwo…
Refractoriness is a fundamental property of excitable elements, such as neurons, indicating the probability for re-excitation in a given time-lag, and is typically linked to the neuronal hyperpolarization following an evoked spike. Here we measured the refractory periods (RPs) in neuronal cultures and observed that an average anisotropic absolute RP could exceed 10 milliseconds and its tail 20 milliseconds, independent of a large stimulation frequency range. It is an order of magnitude longer than anticipated and comparable with the decaying membrane potential timescale. It is followed by a sh…
We develop an extension of the Ott-Antonsen method that allows obtaining the mean activity (spiking rate) of a population of excitable units. By means of the Ott-Antonsen method, equations for the dynamics of the order parameters of coupled excitatory and inhibitory populations of excitable units are obtained, and their mean activities are computed. Two different excitable systems are studied: Adler units and theta neurons. The resulting bifurcation diagrams are compared to those obtained from studying the phenomenological Wilson-Cowan model in some regions of the parameter space. Compatible b…
In neural circuits, statistical connectivity rules strongly depend on neuronal type. Here we study dynamics of neural networks with cell-type specific connectivity by extending the dynamic mean field method, and find that these networks exhibit a phase transition between silent and chaotic activity. By analyzing the locus of this transition, we derive a new result in random matrix theory: the spectral radius of a random connectivity matrix with block-structured variances. We apply our results to show how a small group of hyper-excitable neurons within the network can significantly increase the…
We propose a discrete time dynamical system (a map) as phenomenological model of excitable and spiking-bursting neurons. The model is a discontinuous two-dimensional map. We find condition under which this map has an invariant region on the phase plane, containing chaotic attractor. This attractor creates chaotic spiking-bursting oscillations of the model. We also show various regimes of other neural activities (subthreshold oscillations, phasic spiking etc.) derived from the proposed model.
Experiments suggest that cerebral cortex gains several functional advantages by operating in a dynamical regime near the critical point of a phase transition. However, a long-standing criticism of this hypothesis is that critical dynamics are rather noisy, which might be detrimental to aspects of brain function that require precision. If the cortex does operate near criticality, how might it mitigate the noisy fluctuations? One possibility is that other parts of the brain may act to control the fluctuations and reduce cortical noise. To better understand this possibility, here we numerically a…
Problems with artificial neural networks originate from their deterministic nature and inevitable prior learnings, resulting in inadequate adaptability against unpredictable, abrupt environmental change. Here we show that a stochastically excitable threshold unit can be utilized by these systems to partially overcome the environmental change. Using an excitable threshold system, attractors were created that represent quasi-equilibrium states into which a system settles until disrupted by environmental change. Furthermore, noise-driven attractor stabilization and switching were embodied by inhi…
Many systems are modulated by unknown slow processes. This hinders analysis in highly non-linear systems, such as excitable systems. We show that for such systems, if the input matches the sparse `spiky' nature of the output, the spiking input-output relation can be derived. We use this relation to reproduce and interpret the irregular and complex 1/f response observed in isolated neurons stimulated over days. We decompose the neuronal response into contributions from its long history of internal noise and its short (few minutes) history of inputs, quantifying memory, noise and stability.
Although there is increasing evidence of criticality in the brain, the processes that guide neuronal networks to reach or maintain criticality remain unclear. The present research examines the role of neuronal gain plasticity in time-series of simulated neuronal networks composed of integrate-and-fire stochastic spiking neurons, and the utility of fractal methods in assessing network criticality. Simulated time-series were derived from a network model of fully connected discrete-time stochastic excitable neurons. Monofractal and multifractal analyses were applied to neuronal gain time-series. …
The artificial axon is an excitable node built with the basic biomolecular components and supporting action potentials. Here we demonstrate coincidence firing (the AND operation) and other basic electrophysiology features such as increasing firing rates for increasing input currents. We construct the basic unit for a network by connecting two such excitable nodes through an electronic synapse, producing pre/post synaptic behavior in which one axon induces firing in another. We show that the system is well described by the Hodgkin-Huxley model of nerve excitability, and conclude with a brief ou…
We study some mechanisms responsible for synchronous oscillations and loss of synchrony at physiologically relevant frequencies (10-200 Hz) in a network of heterogeneous inhibitory neurons. We focus on the factors that determine the level of synchrony and frequency of the network response, as well as the effects of mild heterogeneity on network dynamics. With mild heterogeneity, synchrony is never perfect and is relatively fragile. In addition, the effects of inhibition are more complex in mildly heterogeneous networks than in homogeneous ones. In the former, synchrony is broken in two distinc…
The response of a neuron to synaptic input strongly depends on whether or not it has just emitted a spike. We propose a neuron model that after spike emission exhibits a partial response to residual input charges and study its collective network dynamics analytically. We uncover a novel desynchronization mechanism that causes a sequential desynchronization transition: In globally coupled neurons an increase in the strength of the partial response induces a sequence of bifurcations from states with large clusters of synchronously firing neurons, through states with smaller clusters to completel…
Experimental neuroscience increasingly requires tractable models for analyzing and predicting the behavior of neurons and networks. The generalized linear model (GLM) is an increasingly popular statistical framework for analyzing neural data that is flexible, exhibits rich dynamic behavior and is computationally tractable (Paninski, 2004; Pillow et al., 2008; Truccolo et al., 2005). What follows is a brief summary of the primary equations governing the application of GLM's to spike trains with a few sentences linking this work to the larger statistical literature. Latter sections include exten…
Cell assemblies manipulation by optogenetics is pivotal to advance neuroscience and neuroengineering. In in vivo applications, photostimulation often broadly addresses a population of cells simultaneously, leading to feed-forward and to reverberating responses in recurrent microcircuits. The former arise from direct activation of targets downstream, and are straightforward to interpret. The latter are consequence of feedback connectivity and may reflect a variety of time-scales and complex dynamical properties. We investigated wide-field photostimulation in cortical networks in vitro, employin…
To evaluate the nature of the neural code in the cerebral cortex, we have used a combination of theory and experiment to assess how information is represented in a realistic cortical population response. We have shown how a sensory stimulus could be estimated on a biologically-realistic time scale, given brief individual responses from a population of neurons with similar response properties. For neurons in extrastriate motion area MT, a combinatorial code, one that keeps track of the cell identity of action potentials and silences in individual neurons across the population, carries twice as …
To understand how neurons and nervous systems first evolved, we need an account of the origins of neural elongations: Why did neural elongations (axons and dendrites) first originate, such that they could become the central component of both neurons and nervous systems? Two contrasting conceptual accounts provide different answers to this question. Braitenberg's vehicles provide the iconic illustration of the dominant input-output (IO) view. Here the basic role of neural elongations is to connect sensors to effectors, both situated at different positions within the body. For this function, neu…
