For a class of excitatory spiking neuron models with delayed feedback fed with a Poisson stochastic process, it is proven that the stream of output interspike intervals cannot be presented as a Markov process of any order.
Keywords: spiking neuron; Poisson stochastic process; probability density function; delayed feedback; non-Markov stochastic process
Activity of inhibitory neuron with delayed feedback is considered in the framework of point stochastic processes. The neuron receives excitatory input impulses from a Poisson stream, and inhibitory impulses from the feedback line with a delay. We investigate here, how does the presence of inhibitory feedback affect the output firing statistics. Using binding neuron (BN) as a model, we derive analytically the exact expressions for the output interspike intervals (ISI) probability density, mean output ISI and coefficient of variation as functions of model's parameters for the case of threshold 2…
In this paper, we study analytically the impact of an inhibitory autapse on neuronal activity. In order to do this, we formulate conditions on a set of non-adaptive spiking neuron models with delayed feedback inhibition, instead of considering a particular neuronal model. The neuron is stimulated with a stochastic point renewal process of excitatory impulses. Probability density function (PDF) $p(t)$ of output interspike intervals (ISIs) of such a neuron is found exactly without any approximations made. It is expressed in terms of ISIs PDF for the input renewal stream and ISIs PDF for that sam…
For a class of fast {\it Cl-}type inhibitory spiking neuron models with delayed feedback fed with a Poisson stochastic process of excitatory impulses, it is proven that the stream of output interspike intervals cannot be presented as a Markov process of any order.
We present a first-order non-homogeneous Markov model for the interspike-interval density of a continuously stimulated spiking neuron. The model allows the conditional interspike-interval density and the stationary interspike-interval density to be expressed as products of two separate functions, one of which describes only the neuron characteristics, and the other of which describes only the signal characteristics. This allows the use of this model to predict the response when the underlying neuron model is not known or well determined. The approximation shows particularly clearly that signal…
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…
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 the 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 (inter-spike intervals, ISIs) in the network could be non-Markovian. In this paper, we address this question for a sim…
For studying how dynamical responses to external stimuli depend on the synaptic-coupling type, we consider two types of excitatory and inhibitory synchronization (i.e., synchronization via synaptic excitation and inhibition) in complex small-world networks of excitatory regular spiking (RS) pyramidal neurons and inhibitory fast spiking (FS) interneurons. For both cases of excitatory and inhibitory synchronization, effects of synaptic couplings on dynamical responses to external time-periodic stimuli $S(t)$ (applied to a fraction of neurons) are investigated by varying the driving amplitude $A$…
A binding neuron (BN) whith delayed feedback is considered. The neuron is fed externally with a Poisson stream of intensity $λ$. The neuron's output spikes are fed into its input with time delay $Δ$. The resulting output stream of the BN is not Poissonian, and we look for its interspike intervals (ISI) distribution. For BN with threshold 2 an exact mathematical expression as function of $λ$, $Δ$ and BN's internal memory, $τ$ is derived for the ISI distribution, and for higher thresholds it is found numerically. The distributions found are characterized with discontinuities of jump type, and in…
Mathematical modeling is an important tool to study the role of delay in neural systems and to evaluate its effects on the signaling activity of coupled neurons. Models for delayed neurons are often used to represent the dynamics of real neurons, but rarely to assess the energy required to maintain these dynamics. In this work, we address these questions from an energy perspective by considering a pair of Hindmarsh-Rose burst neurons coupled by reciprocal time-delayed coupling with electrical and chemical synapses. We examine the average energy consumption required to maintain cooperative beha…
We consider a single Leaky integrate-and-fire neuron stimulated with Poisson process. We develop a method, which allows one to obtain the first passage time probability density function without any additional approximations.
Using an exactly solvable cortical model of a neuronal network, we show that, by increasing the intensity of shot noise (flow of random spikes bombarding neurons), the network undergoes first- and second-order non-equilibrium phase transitions. We study the nature of the transitions, bursts and avalanches of neuronal activity. Saddle-node and supercritical Hopf bifurcations are the mechanisms of emergence of sustained network oscillations. We show that the network stimulated by shot noise behaves similar to the Morris-Lecar model of a biological neuron stimulated by an applied current.
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 principle, while coupled limit cycle oscillators can overcome mismatch in intrinsic rates and match their frequencies, but zero phase lag synchronization is just achievable in the limit of zero mismatch, i.e., with identical oscillators. Delay in communication, on the other hand, can exert phase shift in the activity of the coupled oscillators. In this study, we address the question of how phase locked, and in particular zero phase lag synchronization, can be achieved for a heterogeneous system of two delayed coupled neurons. We have analytically studied the possibility of inphase synchroni…
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…
Model calculations have been performed on the spike-train response of a pair of Hodgkin-Huxley (HH) neurons coupled by recurrent excitatory-excitatory couplings with time delay. The coupled, excitable HH neurons are assumed to receive the two kinds of spike-train inputs: the transient input consisting of $M$ impulses for the finite duration ($M$: integer) and the sequential input with the constant interspike interval (ISI). The distribution of the output ISI $T_{\rm o}$ shows a rich of variety depending on the coupling strength and the time delay. The comparison is made between the dependence …
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(…
We consider a class of spiking neuronal models, defined by a set of conditions typical for basic threshold-type models, such as the leaky integrate-and-fire or the binding neuron model and also for some artificial neurons. A neuron is fed with a Poisson process. Each output impulse is applied to the neuron itself after a finite delay $Δ$. This impulse acts as being delivered through a fast Cl-type inhibitory synapse. We derive a general relation which allows calculating exactly the probability density function (pdf) $p(t)$ of output interspike intervals of a neuron with feedback based on known…
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 present generalized delayed neural network (DNN) model with positive delay feedback and neuron history. The local stability analysis around trivial local equilibria of delayed neural networks has applied and determine the conditions for the existence of zero root. We develop few innovative delayed neural network models in different dimensions through transformation and extension of some existing models. We found that zero root can have multiplicity two under certain conditions. We further show how the characteristic equation can have zero root and its multiplicity is dependent on the condit…
Probability density function of output interspike intervals is found in exact form for leaky integrate and fire neuron stimulated with Poisson stream. The diffusion approximation is not exploited.
Noise-delayed decay (NDD) phenomenon emerges when the first-spike latency of a periodically forced stochastic neuron exhibits a maximum for a particular range of noise intensity. Here, we investigate the latency response dynamics of a single Hodgkin-Huxley neuron that is subject to both a suprathreshold periodic stimulus and a background activity arriving through dynamic synapses. We study the first spike latency response as a function of the presynaptic firing rate f. This constitutes a more realistic scenario than previous works, since f provides a suitable biophysically realistic parameter …
This paper uses a simple optogenetic model to compare the timing distortion between a randomly-generated target spike sequence and an externally-stimulated neuron spike sequence. Optogenetics is an emerging field of neuroscience where neurons are genetically modified to express light-sensitive receptors that enable external control over when the neurons fire. Given the prominence of neuronal signaling within the brain and throughout the body, optogenetics has significant potential to improve the understanding of the nervous system and to develop treatments for neurological diseases. This paper…
Recent experimental studies strongly suggest the influence of glial purinergic transmission in the modulation of synaptic dynamics. By releasing adenosine triphosphate (ATP), which accumulates as adenosine, astrocytes tonically suppressed synaptic transmission. The delayed multi-step feedback of the glial adenosine with the neuron suggest the existence of hysteresis phenomena, which are investigated in the present study from the theoretical point of view. The model suggests that a memory operator, tripartite synaptic plasticity, governs the mysterious delayed feedback inhibition caused by the …
We describe general characteristics of the Hodgkin-Huxley neuron's response to a periodic train of short current pulses with Gaussian noise. The deterministic neuron is bistable for antiresonant frequencies. When the stimuli arrive at the resonant frequency the firing rate is a continuous function of the current amplitude $I_0$ and scales as $(I_0-I_{th})^{1/2}$, where $I_{th}$ is an approximate threshold. Intervals of continuous irregular response alternate with integer mode-locked regions with bistable excitation edge. There is an even-all multimodal transition between the 2:1 and 3:1 states…
We analyse the storage and retrieval capacity in a recurrent neural network of spiking integrate and fire neurons. In the model we distinguish between a learning mode, during which the synaptic connections change according to a Spike-Timing Dependent Plasticity (STDP) rule, and a recall mode, in which connections strengths are no more plastic. Our findings show the ability of the network to store and recall periodic phase coded patterns a small number of neurons has been stimulated. The self sustained dynamics selectively gives an oscillating spiking activity that matches one of the stored pat…
The inverse Ising model is used in computational neuroscience to infer probability distributions of the synchronous activity of large neuronal populations. This method allows for finding the Boltzmann distribution with single neuron biases and pairwise interactions that maximizes the entropy and reproduces the empirical statistics of the recorded neuronal activity. Here we apply this strategy to large populations of retinal output neurons (ganglion cells) of different types, stimulated by multiple visual stimuli with their own statistics. The activity of retinal output neurons is driven by bot…
Cortical neurons whose activity is recorded in behavioral experiments has been classified into several types such as stimulus-related neurons, delay-period neurons, and reward-related neurons. Moreover, the population activity of neurons during a reaching task can be described by dynamics in low-dimensional space. These results suggest that the low-dimensional dynamics of a few types of neurons emerge in the cerebral cortex that is trained to perform a task. This study investigates a simple neural circuit emerges with a few types of neurons. Assuming an infinite number of neurons in the circui…
Huazhong University of Science and Technology(华中科技大学)
As the third generation of neural networks, spiking neural networks (SNNs) are dedicated to exploring more insightful neural mechanisms to achieve near-biological intelligence. Intuitively, biomimetic mechanisms are crucial to understanding and improving SNNs. For example, the associative long-term potentiation (ALTP) phenomenon suggests that in addition to learning mechanisms between neurons, there are associative effects within neurons. However, most existing methods only focus on the former and lack exploration of the internal association effects. In this paper, we propose a novel Adaptive …
Suppression of excessively synchronous beta-band oscillatory activity in the brain is believed to suppress hypokinetic motor symptoms of Parkinson's disease. Recently, a lot of interest has been devoted to desynchronizing delayed feedback deep brain stimulation (DBS). This type of synchrony control was shown to destabilize the synchronized state in networks of simple model oscillators as well as in networks of coupled model neurons. However, the dynamics of the neural activity in Parkinson's disease exhibits complex intermittent synchronous patterns, far from the idealized synchronous dynamics…
The principles of neural encoding and computations are inherently collective and usually involve large populations of interacting neurons with highly correlated activities. While theories of neural function have long recognized the importance of collective effects in populations of neurons, only in the past two decades has it become possible to record from many cells simulatenously using advanced experimental techniques with single-spike resolution, and to relate these correlations to function and behaviour. This review focuses on the modeling and inference approaches that have been recently d…
Limitations in processing capabilities and memory of today's computers make spiking neuron-based (human) whole-brain simulations inevitably characterized by a compromise between bio-plausibility and computational cost. It translates into brain models composed of a reduced number of neurons and a simplified neuron's mathematical model, leading to the search for new simulation strategies. Taking advantage of the sparse character of brain-like computation, the event-driven technique could represent a way to carry out efficient simulation of large-scale Spiking Neural Networks (SNN). The recent Le…
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…
The effect of intrinsic channel noise is investigated for the dynamic response of a neuronal cell with a delayed feedback loop. The loop is based on the so-called autapse phenomenon in which dendrites establish not only connections to neighboring cells but as well to its own axon. The biophysical modeling is achieved in terms of a stochastic Hodgkin-Huxley model containing such a built in delayed feedback. The fluctuations stem from intrinsic channel noise, being caused by the stochastic nature of the gating dynamics of ion channels. The influence of the delayed stimulus is systematically anal…
We study a network of spiking neurons with heterogeneous excitabilities connected via inhibitory delayed pulses. For globally coupled systems the increase of the inhibitory coupling reduces the number of firing neurons by following a Winner Takes All mechanism. For sufficiently large transmission delay we observe the emergence of collective oscillations in the system beyond a critical coupling value. Heterogeneity promotes neural inactivation and asynchronous dynamics and its effect can be counteracted by considering longer time delays. In sparse networks, inhibition has the counterintuitive e…
Stochastic neural networks are a prototypical computational device able to build a probabilistic representation of an ensemble of external stimuli. Building on the relationship between inference and learning, we derive a synaptic plasticity rule that relies only on delayed activity correlations, and that shows a number of remarkable features. Our "delayed-correlations matching" (DCM) rule satisfies some basic requirements for biological feasibility: finite and noisy afferent signals, Dale's principle and asymmetry of synaptic connections, locality of the weight update computations. Nevertheles…
How spiking neuronal networks encode memories in their different time and spatial scales constitute a fundamental topic in neuroscience and neuro-inspired engineering. Much attention has been paid to large networks and long-term memory, for example in models of associative memory. Smaller circuit motifs may play an important complementary role on shorter time scales, where broader network effects may be of less relevance. Yet, compact computational models of spiking neural networks that exhibit short-term volatile memory and actively hold information until their energy source is switched off, …
We consider here a single-compartment model of these neurons which is capable of describing many of the known features of spike generation, particularly the slow rhythmic pacemaking activity often observed in these cells in a variety of species. Included in the model are ten kinds of voltage dependent ion channels as well as calcium-dependent potassium current. Calcium dynamics includes buffering and pumping. In sections 3-9, each component is considered in detail and parameters estimated from voltage clamp data where possible. In the next two sections simplified versions of some components ar…
Non-invasive methods to measure brain activity are important to understand cognitive processes in the human brain. A prominent example is functional magnetic resonance imaging (fMRI), which is a noisy measurement of a delayed signal that depends non-linearly on the neuronal activity through the neurovascular coupling. These characteristics make the inference of neuronal activity from fMRI a difficult but important step in fMRI studies that require information at the neuronal level. In this article, we address this inference problem using a Bayesian approach where we model the latent neural act…
We analyze a large system of heterogeneous quadratic integrate-and-fire (QIF) neurons with time delayed, all-to-all synaptic coupling. The model is exactly reduced to a system of firing rate equations that is exploited to investigate the existence, stability and bifurcations of fully synchronous, partially synchronous, and incoherent states. In conjunction with this analysis we perform extensive numerical simulations of the original network of QIF neurons, and determine the relation between the macroscopic and microscopic states for partially synchronous states. The results are summarized in t…