Moment-generating function of output stream of leaky integrate-and-fire neuron
Alexander Vidybida, Olha Shchur
备注：
10 pages, 3 figures, manuscript accepted by Ukrainian Journal of Physics
发表信息：
Vidybida, A., & Shchur, O. (2021). Moment-Generating Function of Output Stream of Leaky Integrate-and-Fire Neuron. Ukrainian Journal of Physics, 66(3), 254
The statistics of the output activity of a neuron during its stimulation by the stream of input impulses that forms the stochastic Poisson process is studied. The leaky integrate-and-fire neuron is considered as a neuron model. A new representation of the probability distribution function of the output interspike interval durations is found. Based on it, the moment-generating function of the probability distribution is calculated explicitly. The latter, according to Curtiss theorem, completely determines the distribution itself. In particular, explicit expressions are derived from the moment-generating function for the moments of all orders. The first moment coincides with the one found earlier. Formulas for the second and third moments have been checked numerically by direct modeling of the stochastic dynamics of a neuron with specific physical parameters.
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.
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.
We compute the firing rate of a leaky integrate-and-fire (LIF) neuron with stochastic conductance-based inputs in the limit when synaptic decay times are much shorter than the membrane time constant. A comparison of our analytical results to numeric simulations is presented for a range of biophysically-realistic parameters.
This work reports a transfer function-based approach to characterizing the operation of single neuronal cells in terms of the instantaneous frequency of the input and output signals. We adopt the leaky integrate-and-fire model. The transfer relationship is obtained by performing successive numeric-computational simulations and statistical regressions. Several interesting results are reported, including the identification and characterization of linearity in the transfer relationship, as well as the identification of regions in the parameter space characterized by sharper transfer functions. Th…
The leaky integrate and fire (LIF) neuron represents standard neuronal model used for numerical simulations. The leakage is implemented in the model as exponential decay of trans-membrane voltage towards its resting value. This makes inevitable the usage of machine floating point numbers in the course of simulation. It is known that machine floating point arithmetic is subjected to small inaccuracies, which prevent from exact comparison of floating point quantities. In particular, it is incorrect to decide whether two separate in time states of a simulated system composed of LIF neurons are ex…
The Nonlinear Noisy Leaky Integrate and Fire neuronal models are mathematical models that describe the activity of neural networks. These models have been studied at a microscopic level, using Stochastic Differential Equations, and at a mesoscopic/macroscopic level, through the mean field limits using Fokker-Planck type equations. The aim of this paper is to improve their understanding, using a numerical study of their particle systems. We analyse in depth the behaviour of the classical and physical solutions of the Stochastic Differential Equations and, we compare it with what is already know…
The first-passage-time distribution of a leaky integrate-and-fire neuron driven by a characteristically coloured noise is approximated by matching a transient and a steady-state solution of the membrane voltage distribution. These approximations follow from a simple manipulation, made possible by the specific `eigen' colouring of the noise, which allows to express the membrane potential as a Gaussian diffusion process on top of a deterministic exponential movement. Following, the presented method is extended to the case of an arbitrarily coloured noise driving by factoring out the `eigen' nois…
Providing an analytical treatment to the stochastic feature of neurons' dynamics is one of the current biggest challenges in mathematical biology. The noisy leaky integrate-and-fire model and its associated Fokker-Planck equation are probably the most popular way to deal with neural variability. Another well-known formalism is the escape-rate model: a model giving the probability that a neuron fires at a certain time knowing the time elapsed since its last action potential. This model leads to a so-called age-structured system, a partial differential equation with non-local boundary condition …
We present a biophysical approach for the coupling of neural network activity as resulting from proper dipole currents of cortical pyramidal neurons to the electric field in extracellular fluid. Starting from a reduced threecompartment model of a single pyramidal neuron, we derive an observation model for dendritic dipole currents in extracellular space and thereby for the dendritic field potential that contributes to the local field potential of a neural population. This work aligns and satisfies the widespread dipole assumption that is motivated by the "open-field" configuration of the dendr…
We study the collective dynamics of a Leaky Integrate and Fire network in which precise relative phase relationship of spikes among neurons are stored, as attractors of the dynamics, and selectively replayed at differentctime scales. Using an STDP-based learning process, we store in the connectivity several phase-coded spike patterns, and we find that, depending on the excitability of the network, different working regimes are possible, with transient or persistent replay activity induced by a brief signal. We introduce an order parameter to evaluate the similarity between stored and recalled …
Artificial neural networks (ANNs) have been extensively used for the description of problems arising from biological systems and for constructing neuromorphic computing models. The third generation of ANNs, namely, spiking neural networks (SNNs), inspired by biological neurons enable a more realistic mimicry of the human brain. A large class of the problems from these domains is characterized by the necessity to deal with the combination of neurons, spikes and synapses via integrate-and-fire neuron models. Motivated by important applications of the integrate-and-fire of neurons in neuromorphic…
We study the synchronization of fully-connected and totally excitatory integrate and fire neural networks in presence of Gaussian white noises. Using a large deviation principle, we prove the stability of the synchronized state under stochastic perturbations. Then, we give a lower bound on the probability of synchronization for networks which are not initially synchronized. This bound shows the robustness of the emergence of synchronization in presence of small stochastic perturbations.
We study the dynamics of identical leaky integrate-and-fire neurons with symmetric non-local coupling. Upon varying control parameters (coupling strength, coupling range, refractory period) we investigate the system's behaviour and highlight the formation of chimera states. We show that the introduction of a refractory period enlarges the parameter region where chimera states appear and affects the chimera multiplicity.
We show that the stochastic Morris-Lecar neuron, in a neighborhood of its stable point, can be approximated by a two-dimensional Ornstein-Uhlenbeck (OU) modulation of a constant circular motion. The associated radial OU process is an example of a leaky integrate-and-fire (LIF) model prior to firing. A new model constructed from a radial OU process together with a simple firing mechanism based on detailed Morris-Lecar firing statistics reproduces the Morris-Lecar Interspike Interval (ISI) distribution, and has the computational advantages of a LIF. The result justifies the large amount of atten…
The response properties of excitable systems driven by colored noise are of great interest, but are usually mathematically only accessible via approximations. For this reason, dichotomous noise, a rare example of a colored noise leading often to analytically tractable problems, has been extensively used in the study of stochastic systems. Here, we calculate exact expressions for the power spectrum and the susceptibility of a leaky integrate-and-fire neuron driven by asymmetric dichotomous noise. While our results are in excellent agreement with simulations, they also highlight a limitation of …
One of the most important challenges in mathematical neuroscience is to properly illustrate the stochastic nature of neurons. Among different approaches, the noisy leaky integrate-and-fire and the escape rate models are probably the most popular. These two models are usually chosen to express different noise action over the neural cell. In this paper we investigate the link between the two formalisms in the case of a neuron subject to a time dependent input. To this aim, we introduce a new general stochastic framework. As we shall prove, our general framework entails the two already existing o…
We derive analytical formulae for the firing rate of integrate-and-fire neurons endowed with realistic synaptic dynamics. In particular we include the possibility of multiple synaptic inputs as well as the effect of an absolute refractory period into the description.
Statistical properties of spike trains as well as other neurophysiological data suggest a number of mathematical models of neurons. These models range from entirely descriptive ones to those deduced from the properties of the real neurons. One of them, the diffusion leaky integrate-and-fire neuronal model, which is based on the Ornstein-Uhlenbeck stochastic process that is restricted by an absorbing barrier, can describe a wide range of neuronal activity in terms of its parameters. These parameters are readily associated with known physiological mechanisms. The other model is descriptive, Gamm…
We investigate numerically the collective dynamical behavior of pulse-coupled non-leaky integrate-and-fire-neurons that are arranged on a two-dimensional small-world network. To ensure ongoing activity, we impose a probability for spontaneous firing for each neuron. We study network dynamics evolving from different sets of initial conditions in dependence on coupling strength and rewiring probability. Beside a homogeneous equilibrium state for low coupling strength, we observe different local patterns including cyclic waves, spiral waves, and turbulent-like patterns, which -- depending on netw…
Neural network dynamics emerge from the interaction of spiking cells. One way to formulate the problem is through a theoretical framework inspired by ideas coming from statistical physics, the so-called mean-field theory. In this document, we investigate different issues related to the mean-field description of an excitatory network made up of leaky integrate-and-fire neurons. The description is written in the form a nonlinear partial differential equation which is known to blow up in finite time when the network is strongly connected. We prove that in a moderate coupling regime the equation i…
We extensively explore networks of weakly unbalanced, leaky integrate-and-fire (LIF) neurons for different coupling strength, connectivity, and by varying the degree of refractoriness, as well as the delay in the spike transmission. We find that the neural network does not only exhibit a microscopic (single-neuron) stochastic-like evolution, but also a collective irregular dynamics (CID). Our analysis is based on the computation of a suitable order parameter, typically used to characterize synchronization phenomena and on a detailed scaling analysis (i.e. simulations of different network sizes…
Tactile afferents such as (RA), and Pacinian (PC) afferents that respond to external stimuli enable complicated actions such as grasping, stroking and identifying an object. To understand the tactile sensation induced by these actions deeply, the activities of the tactile afferents need to be revealed. For this purpose, we develop a computational model for each tactile afferent for vibration stimuli, combining finite element analysis finite element method (FEM) analysis and a leaky integrate-and-fire model that represents the neural characteristics. This computational model can easily estimate…
Several experimental studies show the existence of leader neurons in population bursts of 2D living neural networks. A leader neuron is, basically, a neuron which fires at the beginning of a burst (respectively network spike) more often that we expect by looking at its whole mean neural activity. This means that leader neurons have some burst triggering power beyond a simple statistical effect. In this study, we characterize these leader neuron properties. This naturally leads us to simulate neural 2D networks. To build our simulations, we choose the leaky integrate and fire (lIF) neuron model…
Author summary: Synchronization of neuronal spiking in the brain is related to cognitive functions, such as perception, attention, and memory. It is therefore important to determine which properties of neurons influence their collective behavior in a network and to understand how. A prominent feature of many cortical neurons is spike frequency adaptation, which is caused by slow transmembrane currents. We investigated how these adaptation currents affect the synchronization tendency of coupled model neurons. Using the efficient adaptive exponential integrate-and-fire (aEIF) model and a biophys…
We studied the impact of a dynamical threshold on the f-I curve-the relationship between the input and the firing rate of a neuron-in the presence of background synaptic inputs. First, we found that, while the leaky integrate-and-fire model cannot reproduce the f-I curve of a cortical neuron, the leaky integrate-and-fire model with dynamical threshold can reproduce it very well. Second, we found that the dynamical threshold modulates the onset and the asymptotic behavior of the f-I curve. These results suggest that a cortical neuron has an adaptation mechanism and that the dynamical threshold …
Reduced models of neuronal activity such as Integrate-and-Fire models allow a description of neuronal dynamics in simple, intuitive terms and are easy to simulate numerically. We present a method to fit an Integrate-and-Fire-type model of neuronal activity, namely a modified version of the Spike Response Model, to a detailed Hodgkin-Huxley-type neuron model driven by stochastic spike arrival. In the Hogkin-Huxley model, spike arrival at the synapse is modeled by a change of synaptic conductance. For such conductance spike input, more than 70% of the postsynaptic action potentials can be predic…
The collective dynamics of neural populations are often characterized in terms of correlations in the spike activity of different neurons. Open questions surround the basic nature of these correlations. In particular, what leads to higher-order correlations -- correlations in the population activity that extend beyond those expected from cell pairs? Here, we examine this question for a simple, but ubiquitous, circuit feature: common fluctuating input arriving to spiking neurons of integrate-and-fire type. We show that leads to strong higher-order correlations, as for earlier work with discrete…
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. …
In this paper, we propose a shot noise-based leaky integrated and firing neuron model and provide a detailed analysis of the performance of this model compared to the traditional diffusion approximated model. In theoretical neuroscience, there are three general neuron models in the field: Compartmental neuron model is a conductance-based model, in which it views the biological neurons as a large circuit. The problem of this model comes from its structural complexity and the number of its free parameters; Leaky integrated and firing model is a more flexible model due to the special design calle…
How extracellular electric fields, as generated endogenously or through transcranial brain stimulation, affect the dynamics of large neuronal populations is of great interest but not well understood. To study the collective dynamics of large populations single-compartment (point) model neurons have been proven very successful. These models, however, lack the dendritic morphology to biophysically account for the effects of electric fields, and for changes in synaptic integration due to morphology alone. Here we (i) characterize the response of a canonical spatial (ball-and-stick) model neuron t…
Identifying the brain's neuronal cluster size to be presented as nodes in a network computation is critical to both neuroscience and artificial intelligence, as these define the cognitive blocks required for building intelligent computation. Experiments support many forms and sizes of neural clustering, while neural mass models (NMM) assume scale-invariant functionality. Here, we use computational simulations with brain-derived fMRI network to show that not only brain network stays structurally self-similar continuously across scales, but also neuron-like signal integration functionality is pr…
Friedrich Alexander University Erlangen-Nürnberg (FAU)（埃尔朗根-纽伦堡弗里德里希-亚历山大大学（FAU））；University Hospital Erlangen（埃尔朗根大学医院）；Friedrich-Alexander University Erlangen-Nürnberg (FAU)（埃尔朗根-纽伦堡弗里德里希-亚历山大大学（FAU））
Up to now, modern Machine Learning is mainly based on fitting high dimensional functions to enormous data sets, taking advantage of huge hardware resources. We show that biologically inspired neuron models such as the Leaky-Integrate-and-Fire (LIF) neurons provide novel and efficient ways of information encoding. They can be integrated in Machine Learning models, and are a potential target to improve Machine Learning performance. Thus, we derived simple update-rules for the LIF units from the differential equations, which are easy to numerically integrate. We apply a novel approach to train th…
Voltage-sensitive dye imaging (VSDi) has revealed fundamental properties of neocortical processing at mesoscopic scales. Since VSDi signals report the average membrane potential, it seems natural to use a mean-field formalism to model such signals. Here, we investigate a mean-field model of networks of Adaptive Exponential (AdEx) integrate-and-fire neurons, with conductance-based synaptic interactions. The AdEx model can capture the spiking response of different cell types, such as regular-spiking (RS) excitatory neurons and fast-spiking (FS) inhibitory neurons. We use a Master Equation formal…
When stimulated by a very large number of Poisson-like presynaptic current input spikes, the temporal dynamics of the soma membrane potential $V(t)$ of a leaky integrate-and-fire (LIF) neuron is typically modeled in the diffusion limit and treated as a Ornstein-Uhlenbeck process (OUP). When the potential reaches a threshold value $θ$, $V(t) = θ$, the LIF neuron fires and the membrane potential is reset to a resting value, $V_0 < θ$, and clamped to this value for a specified (non-stochastic) absolute refractory period $T_r \ge 0$, after which the cycle is repeated. The time between firings is g…
We propose another integrate-and-fire model as a single neuron model. We study a globally coupled noisy integrate-and-fire model with inhibitory interaction using the Fokker-Planck equation and the Langevin equation, and find a reentrant transition of oscillatory states. Intermittent time evolutions of neuron firing are found in strongly inhibited systems. We propose another integrate-and-fire-or-burst model including the dynamics of the low-threshold Ca$^{2+}$ current based on the new integrate-and-fire model. We study a globally coupled noisy integrate-and-fire-or-burst model with inhibitory…
The spiking activity of single neurons can be well described by a nonlinear integrate-and-fire model that includes somatic adaptation. When exposed to fluctuating inputs sparsely coupled populations of these model neurons exhibit stochastic collective dynamics that can be effectively characterized using the Fokker-Planck equation. [...] Here we derive from that description four simple models for the spike rate dynamics in terms of low-dimensional ordinary differential equations using two different reduction techniques: one uses the spectral decomposition of the Fokker-Planck operator, the othe…
We consider a threshold-crossing spiking process as a simple model for the activity within a population of neurons. Assuming that these neurons are driven by a common fluctuating input with Gaussian statistics, we evaluate the cross-correlation of spike trains in pairs of model neurons with different thresholds. This correlation function tends to be asymmetric in time, indicating a preference for the neuron with the lower threshold to fire before the one with the higher threshold, even if their inputs are identical. The relationship between these results and spike statistics in other models of…
The Expressive Leaky Memory Neuron: an Efficient and Expressive Phenomenological Neuron Model Can Solve Long-Horizon Tasks
University of Tübingen（蒂宾根大学）；Max Planck Institute for Intelligent Systems（马克斯·普朗克智能系统研究所）；Mila, Quebec AI Institute（米拉魁北克人工智能研究所）；Max Planck Institute for Biological Cybernetics（马克斯·普朗克生物控制论研究所）
Variability in neural responses is an ubiquitous phenomenon in neurons, usually modeled with stochastic differential equations. In particular, stochastic integrate-and-fire models are widely used to simplify theoretical studies. The statistical properties of the generated spikes depend on the stimulating input current. Given that real sensory neurons are driven by time-dependent signals, here we study how the inter-spike interval distribution of integrate-and-fire neurons depends on the evolution of the stimulus, in a quasi-static limit. We obtain a closed-form expression for this distribution…
Spike-train responses of single Hodgkin-Huxley (HH) and integrate-and-fire (IF) neurons with and without the refractory period, are calculated and compared. The HH and IF neurons are assumed to receive spike-train inputs with the constant interspike intervals (ISIs) and stochastic ISIs given by the Gamma distribution, through excitatory and inhibitory synaptic couplings: for both the couplings the HH neuron can fire while the IF neuron can only for the excitatory one. It is shown that the response to the constant-ISI inputs of the IF neuron strongly depends on the refractory period and the syn…
Stochastic integrate-and-fire (IF) neuron models have found widespread applications in computational neuroscience. Here we present results on the white-noise-driven perfect, leaky, and quadratic IF models, focusing on the spectral statistics (power spectra, cross spectra, and coherence functions) in different dynamical regimes (noise-induced and tonic firing regimes with low or moderate noise). We make the models comparable by tuning parameters such that the mean value and the coefficient of variation of the interspike interval match for all of them. We find that, under these conditions, the p…
Qualitatively, some real networks in the brain could be characterized as 'small worlds', in the sense that the structure of their connections is intermediate between the extremes of an orderly geometric arrangement and of a geometry-independent random mesh. Small worlds can be defined more precisely in terms of their mean path length and clustering coefficient; but is such a precise description useful to better understand how the type of connectivity affects memory retrieval? We have simulated an autoassociative memory network of integrate-and-fire units, positioned on a ring, with the network…
Nonlinear Noisy Leaky Integrate and Fire (NNLIF) models for neurons networks can be written as Fokker-Planck-Kolmogorov equations on the probability density of neurons, the main parameters in the model being the connectivity of the network and the noise. We analyse several aspects of the NNLIF model: the number of steady states, a priori estimates, blow-up issues and convergence toward equilibrium in the linear case. In particular, for excitatory networks, blow-up always occurs for initial data concentrated close to the firing potential. These results show how critical is the balance between n…
Randomly-connected networks of integrate-and-fire (IF) neurons are known to display asynchronous irregular (AI) activity states, which resemble the discharge activity recorded in the cerebral cortex of awake animals. However, it is not clear whether such activity states are specific to simple IF models, or if they also exist in networks where neurons are endowed with complex intrinsic properties similar to electrophysiological measurements. Here, we investigate the occurrence of AI states in networks of nonlinear IF neurons, such as the adaptive exponential IF (Brette-Gerstner-Izhikevich) mode…
ILIF: Temporal Inhibitory Leaky Integrate-and-Fire Neuron for Overactivation in Spiking Neural Networks
Department of Data Science and AI, Monash University（墨尔本大学数据科学与人工智能系）；School of Computer Science and Engineering, Southeast University（东南大学计算机科学与工程学院）；School of Software, Northeastern University（东北大学软件学院）
The binding neuron model is inspired by numerical simulation of Hodgkin-Huxley-type point neuron, as well as by the leaky integrate-and-fire model. In the binding neuron, the trace of an input is remembered for a fixed period of time after which it disappears completely. This is in the contrast with the above two models, where the postsynaptic potentials decay exponentially and can be forgotten only after triggering. The finiteness of memory in the binding neuron allows one to construct fast recurrent networks for computer modeling. Recently, the finiteness is utilized for exact mathematical d…
Computational models of the respiratory central pattern generator (rCPG) are usually based on biologically-plausible Hodgkin Huxley neuron models. Such models require numerous parameters and thus are prone to overfitting. The HH approach is motivated by the assumption that the biophysical properties of neurons determine the network dynamics. Here, we implement the rCPG using simpler Izhikevich resonate-and-fire neurons. Our rCPG model generates a 3-phase respiratory motor pattern based on established connectivities and can reproduce previous experimental and theoretical observations. Further, …
By controlling the state of neuronal populations, neuromodulators ultimately affect behaviour. A key neuromodulation mechanism is the alteration of neuronal excitability via the modulation of ion channel expression. This type of neuromodulation is normally studied via conductance-based models, but those models are computationally challenging for large-scale network simulations needed in population studies. This paper studies the modulation properties of the Multi-Quadratic Integrate-and-Fire (MQIF) model, a generalisation of the classical Quadratic Integrate-and-Fire (QIF) model. The model is …
One of the fundamental characteristics of a nonlinear system is how it transfers correlations in its inputs to correlations in its outputs. This is particularly important in the nervous system, where correlations between spiking neurons are prominent. Using linear response and asymptotic methods for pairs of unconnected integrate-and-fire (IF) neurons receiving white noise inputs, we show that this correlation transfer depends on the output spike firing rate in a strong, stereotyped manner, and is, surprisingly, almost independent of the interspike variance. For cells receiving heterogeneous i…
Noise in spiking neurons is commonly modeled by a noisy input current or by generating output spikes stochastically with a voltage-dependent hazard rate ("escape noise"). While input noise lends itself to modeling biophysical noise processes, the phenomenological escape noise is mathematically more tractable. Using the level-crossing theory for differentiable Gaussian processes, we derive an approximate mapping between colored input noise and escape noise in leaky integrate-and-fire neurons. This mapping requires the first-passage-time (FPT) density of an overdamped Brownian particle driven by…
The task of the brain is to look for structure in the external input. We study a network of integrate-and-fire neurons with several types of recurrent connections that learns the structure of its time-varying feedforward input by attempting to efficiently represent this input with spikes. The efficiency of the representation arises from incorporating the structure of the input into the decoder, which is implicit in the learned synaptic connectivity of the network. While in the original work of [Boerlin, Machens, Denève 2013] and [Brendel et al., 2017] the structure learned by the network to ma…
We explore the dynamics of an integrate-and-fire neuron with an oscillatory stimulus. The frustration due to the competition between the neuron's natural firing period and that of the oscillatory rhythm, leads to a rich structure of asymptotic phase locking patterns and ordering dynamics. The phase transitions between these states can be classified as either tangent or discontinuous bifurcations, each with its own characteristic scaling laws. The discontinuous bifurcations exhibit a new kind of phase transition that may be viewed as intermediate between continuous and first order, while tangen…
