An integral equation describing the time evolution of the population activity in a homogeneous pool of spiking neurons of the integrate-and-fire type is discussed. It is analytically shown that transients from a state of incoherent firing can be immediate. The stability of incoherent firing is analyzed in terms of the noise level and transmission delay and a bifurcation diagram is derived. The response of a population of noisy integrate-and-fire neurons to an input current of small amplitude is calculated and characterized by a linear filter L. The stability of perfectly synchronized `locked' solutions is analyzed.

We study analytically the dynamics of a network of sparsely connected inhibitory integrate-and-fire neurons in a regime where individual neurons emit spikes irregularly and at a low rate. In the limit when the number of neurons N → ∞, the network exhibits a sharp transition between a stationary and an oscillatory global activity regime where neurons are weakly synchronized. The activity becomes oscillatory when the inhibitory feedback is strong enough. The period of the global oscillation is found to be mainly controlled by synaptic times, but depends also on the characteristics of the external input. In large but finite networks, the analysis shows that global oscillations of finite coherence time generically exist both above and below the critical inhibition threshold. Their characteristics are determined as functions of systems parameters, in these two different regimes. The results are found to be in good agreement with numerical simulations.

It is generally believed that a neuron is a threshold element which fires when some variable u reaches a threshold. Here we pursue the question of whether this picture can be justified and study the four-dimensional neuron model of Hodgkin and Huxley as a concrete example. The model is approximated by a response kernel expansion in terms of a single variable, the membrane voltage. The first-order term is linear in the input and has the typical form of an elementary postsynaptic potential. Higher-order kernels take care of nonlinear interactions between input spikes. In contrast to the standard Volterra expansion the kernels depend on the firing time of the most recent output spike. In particular, a zero-order kernel which describes the shape of the spike and the typical afterpotential is included. Our model neuron fires, if the membrane voltage, given by the truncated response kernel expansion crosses a threshold. The threshold model is tested on a spike train generated by t...

The nature and origin of the temporal irregularity in the electrical activity of cortical neurons in vivo are still not well understood. We consider the hypothesis that this irregularity is due to a balance of excitatory and inhibitory currents into the cortical cells. We study a network model with excitatory and inhibitory populations of simple binary units. The internal feedback is mediated by relatively large synaptic strengths, so that the magnitude of the total excitatory as well as inhibitory feedback is much larger than the neuronal threshold. The connectivity is random and sparse. The mean number of connections per unit is large but small compared to the total number of cells in the network. The network also receives a large, temporally regular input from external sources. An analytical solution of the mean-field theory of this model which is exact in the limit of large network size is presented. This theory reveals a new cooperative stationary state of large networks, which we term a balanced state. In this state, a balance between the excitatory and inhibitory inputs emerges dynamically for a wide range of parameters, resulting in a net input whose temporal fluctuations are of the same order as its mean. The internal synaptic inputs act as a strong negative feedback, which linearizes the population responses to the external drive despite the strong nonlinearity of the individual cells. This feedback also greatly stabilizes 1 the system's state and enables it to track a time-dependent input on time scales much shorter than the time constant of a single cell. The spatio-temporal statistics of the balanced state is calculated. It is shown that the auto-correlations decay on a short time scale yielding an approximate Poissonian temporal s...

We demonstrate that single-variable integrate-and-fire models can quantitatively capture the dynamics of a physiologically-detailed model for fast-spiking cortical neurons. Through a systematic set of approximations, we reduce the conductance based model to two variants of integrate-and-fire models. In the first variant (non-linear integrate-and-fire model), parameters depend on the instantaneous membrane potential whereas in the second variant, they depend on the time elapsed since the last spike (Spike Response Model). The direct reduction links features of the simple models to biophysical features of the full conductance based model. To quantitatively

Present and permanent address: Physik-Department der TU Munchen Exploiting local stability we show what neuronal characteristics are essential to ensure that coherent oscillations are asymptotically stable in a spatially homogeneous network of spiking neurons. Under standard conditions, a necessary and in the limit of a large number of interacting neighbors also sufficient condition is that the postsynaptic potential is increasing in time as the neurons fire. If the postsynaptic potential is decreasing, oscillations are bound to be unstable. This is a kind of locking theorem and boils down to a subtle interplay of axonal delays, postsynaptic potentials, and refractory behavior. The theorem also allows for mixtures of excitatory and inhibitory interactions. On the basis of the locking theorem we present a simple geometric method to verify existence and local stability of a coherent oscillation. 2 1

Introduction Neuronal activity due to recurrent excitations in the form of a spatially localized pulse or bump has been proposed as a mechanism for feature selectivity in models of the visual system (Somers, Nelson, & Sur, 1995; Hansel & Sompolinsky, 1998), the head direction system (Skaggs, Knieram, Kudrimoti, & McNaughton, 1995; Zhang, 1996; Redish, Elga, & Touretzky, 1996), and working memory (Wilson & Cowan, 1973; Amit & Brunel, 1997; Camperi & Wang, 1998). Many of the previous mathematical formulations of such structures have employedpopulation rate models (Wilson &Cowan, 1972, 1973; Amari, 1977; Kishimoto & Amari, 1979; Hansel & Sompolinsky, 1998). (See Ermentrout, 1998, for a recent review.) Here, we consider a network of spiking neurons that shows such structures and investigate their properties. In our network we #nd localized time-stationary states

The prevalence of coherent oscillations in various frequency ranges in the central nervous system raises the question of the mechanisms that synchronize large populations of neurons. We study synchronization in models of large networks of spiking neurons with random sparse connectivity. Synchrony occurs only when the average number of synapses, M , that a cell receives is larger than a critical value, M c . Below M c , the system is in an asynchronous state. In the limit of weak coupling, assuming identical neurons, we reduce the model to a system of phase oscillators which are coupled via an effective interaction, \Gamma. In this framework, we develop an approximate theory for sparse networks of identical neurons to estimate M c analytically from the Fourier coefficients of \Gamma. Our approach relies on the assumption that the dynamics of a neuron depend mainly on the number of cells that are presynaptic to it. We apply this theory to compute M c for a model of inhibitory networks of integrate-and-fire (I&F) neurons as a function of the intrinsic neuronal properties (e.g., the refractory period T r ), the synaptic time constants and the strength of the external stimulus, I ext . The number M c is found to be non-monotonous with the strength of I ext . For T r = 0, we estimate the minimum value of M c over all the parameters of the model to be 363:8. Above M c , the neurons tend to fire in: 1) smeared one cluster states at high firing rates and 2) smeared two or more cluster states at low firing rates. Refractoriness decreases M c at intermediate and high firing rates. These results are compared against numerical simulations. We show numerically that systems with different sizes, N , behave in the same way provided the connectivity, M , is such a way that 1=M eff = 1=...

When the local field potential of a cortical network displays coherent fast oscillations (~40-Hz gamma or ~200-Hz sharp-wave ripples), the spike trains of constituent neurons are typically irregular and sparse. The dichotomy between rhythmic local field and stochastic spike trains presents a challenge to the theory of brain rhythms in the framework of coupled oscillators. Previous studies have shown that when noise is large and recurrent inhibition is strong, a coherent network rhythm can be generated while single neurons fire intermittently at low rates compared to the frequency of the oscillation. However, these studies used too simplified synaptic kinetics to allow quantitative predictions of the population rhythmic frequency. Here we show how to derive quantitatively the coherent

We calculate the firing rate of the quadratic integrate-and-fire neuron in response to a colored noise input current. Such an input current is a good approximation to the noise due to the random bombardment of spikes, with the correlation time of the noise corresponding to the decay time of the synapses. The key parameter that determines the firing rate is the ratio of the correlation time of the colored noise, ¿s, to the neuronal time constant, ¿m. We calculate the firing rate exactly in two limits: when the ratio, ¿s=¿m, goes to zero (white noise) and when it goes to infinity. The correction to the short correlation time limit is O.¿s=¿m/, which is qualitatively different from that of the leaky integrate-and-fire neuron, where the correction is O. p ¿s=¿m/. The difference is due to the different boundary conditions of the probability density function of the membrane potential of the neuron at firing threshold. The correction to the long correlation time limit is O.¿m=¿s/. By combining the short and long correlation time limits, we derive an expression that provides a good approximation to the firing rate over the whole range of ¿s=¿m in the suprathreshold regime— that is, in a regime in which the average current is sufficient to make the cell fire. In the subthreshold regime, the expression breaks down somewhat when ¿s becomes large compared to ¿m.