The computation performed by a neuron can be formulated as a combination of dimensional reduction in stimulus space and the nonlinearity inherent in a spiking output. White noise stimulus and reverse correlation (the spike-triggered average and spike-triggered covariance) are often used in experimental neuroscience to `ask' neurons which dimensions in stimulus space they are sensitive to, and to characterize the nonlinearity of the response. In this paper, we apply reverse correlation to the simplest model neuron with temporal dynamics--the leaky integrate-and-fire model--and find that even for this simple case standard techniques do not recover the known neural computation. To overcome this, we develop novel reverse correlation techniques by selectively analyzing only `isolated' spikes, and taking explicit account of the extended silences that precede these isolated spikes. We discuss the implications of our methods to the characterization of neural adaptation. Although these methods are developed in the context of the leaky integrate-and-fire model, our findings are relevant for the analysis of spike trains from real neurons.
Spike correlations between neurons are ubiquitous in the cortex, but their role is at present not understood. Here we describe the firing response of a leaky integrate-and-fire neuron (LIF) when it receives a temporarily correlated input generated by presynaptic correlated neuronal populations. Input correlations are characterized in terms of the firing rates, Fano factors, correlation coefficients and correlation timescale of the neurons driving the target neuron. We show that the sum of the presynaptic spike trains cannot be well described by a Poisson process. Solutions of the output firing…
Spike generation in neurons produces a temporal point process, whose statistics is governed by intrinsic phenomena and the external incoming inputs to be coded. In particular, spike-evoked adaptation currents support a slow temporal process that conditions spiking probability at the present time according to past activity. In this work, we study the statistics of interspike interval correlations arising in such non-renewal spike trains, for a neuron model that reproduces different spike modes in a small adaptation scenario. We found that correlations are stronger as the neuron fires at a parti…
An analytical description of the response properties of simple but realistic neuron models in the presence of noise is still lacking. We determine completely up to the second order the firing statistics of a single and a pair of leaky integrate-and-fire neurons (LIFs) receiving some common slowly filtered white noise. In particular, the auto- and cross-correlation functions of the output spike trains of pairs of cells are obtained from an improvement of the adiabatic approximation introduced in \cite{Mor+04}. These two functions define the firing variability and firing synchronization between …
We are interested in characterization of population synchronization of bursting neurons which exhibit both the slow bursting and the fast spiking timescales, in contrast to spiking neurons. Population synchronization may be well visualized in the raster plot of neural spikes which can be obtained in experiments. The instantaneous population firing rate (IPFR) $R(t)$, which may be directly obtained from the raster plot of spikes, is often used as a realistic collective quantity describing population behaviors in both the computational and the experimental neuroscience. For the case of spiking n…
We are interested in characterization of synchronization transitions of bursting neurons in the frequency domain. Instantaneous population firing rate (IPFR) $R(t)$, which is directly obtained from the raster plot of neural spikes, is often used as a realistic collective quantity describing population activities in both the computational and the experimental neuroscience. For the case of spiking neurons, a realistic time-domain order parameter, based on $R(t)$, was introduced in our recent work to characterize the spike synchronization transition. Unlike the case of spiking neurons, the IPFR $…
This paper predicts the ability to externally control the firing times of a cortical neuron whose behavior follows the Izhikevich neuron model. The Izhikevich neuron model provides an efficient and biologically plausible method to track a cortical neuron's membrane potential and its firing times. The external control is a simple optogenetic model represented by a constant current source that can be turned on or off. This paper considers a firing frequency that is sufficiently low for the membrane potential to return to its resting potential after it fires. The time required for the neuron to c…
For modeling complex synaptic connectivity, we consider the Watts-Strogatz small-world network which interpolates between regular lattice and random network via rewiring, and investigate the effect of small-world connectivity on emergence of noise-induced population synchronization in an inhibitory population of subthreshold bursting Hindmarsh-Rose neurons. Thus, noise-induced slow burst synchronization and fasg spike synchronization are found to appear in a synchronized region of the $J-D$ plane. As the rewiring probability $p$ is decreased from 1 (random network) to 0 (regular lattice), the …
This paper predicts the ability to externally control the firing times of a cortical neuron whose behavior follows the Izhikevich neuron model. The Izhikevich neuron model provides an efficient and biologically plausible method to track a cortical neuron's membrane potential and its firing times. The external control is a simple optogenetic model represented by an illumination source that stimulates a saturating or decaying membrane current. This paper considers firing frequencies that are sufficiently low for the membrane potential to return to its resting potential after it fires. The time r…
We observe and study a self-organized phenomenon whereby the activity in a network of spiking neurons spontaneously terminates. We consider different types of populations, consisting of bistable model neurons connected electrically by gap junctions, or by either excitatory or inhibitory synapses, in a scale-free connection topology. We find that strongly synchronized population spiking events lead to complete cessation of activity in excitatory networks, but not in gap junction or inhibitory networks. We identify the underlying mechanism responsible for this phenomenon by examining the particu…
We demonstrate the efficacy of a new spike-sorting method based on a Markov Chain Monte Carlo (MCMC) algorithm by applying it to real data recorded from Purkinje cells (PCs) in young rat cerebellar slices. This algorithm is unique in its capability to estimate and make use of the firing statistics as well as the spike amplitude dynamics of the recorded neurons. PCs exhibit multiple discharge states, giving rise to multimodal interspike interval (ISI) histograms and to correlations between successive ISIs. The amplitude of the spikes generated by a PC in an "active" state decreases, a feature t…
Brain plasticity, also known as neuroplasticity, is a fundamental mechanism of neuronal adaptation in response to changes in the environment or due to brain injury. In this review, we show our results about the effects of synaptic plasticity on neuronal networks composed by Hodgkin-Huxley neurons. We show that the final topology of the evolved network depends crucially on the ratio between the strengths of the inhibitory and excitatory synapses. Excitation of the same order of inhibition revels an evolved network that presents the rich-club phenomenon, well known to exist in the brain. For ini…
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 …
We show that in model neuronal cultures, where the probability of interneuronal connection formation decreases exponentially with increasing distance between the neurons, there exists a small number of spatial nucleation centers of a network spike, from where the synchronous spiking activity starts propagating in the network typically in the form of circular traveling waves. The number of nucleation centers and their spatial locations are unique and unchanged for a given realization of neuronal network but are different for different networks. In contrast, if the probability of interneuronal c…
Spiking neuronal networks are usually simulated with three main simulation schemes: the classical time-driven and event-driven schemes, and the more recent hybrid scheme. All three schemes evolve the state of a neuron through a series of checkpoints: equally spaced in the first scheme and determined neuron-wise by spike events in the latter two. The time-driven and the hybrid scheme determine whether the membrane potential of a neuron crosses a threshold at the end of of the time interval between consecutive checkpoints. Threshold crossing can, however, occur within the interval even if this t…
In this article, our wish is to demystify some aspects of coding with spike-timing, through a simple review of well-understood technical facts regarding spike coding. The goal is to help better understanding to which extend computing and modelling with spiking neuron networks can be biologically plausible and computationally efficient. We intentionally restrict ourselves to a deterministic dynamics, in this review, and we consider that the dynamics of the network is defined by a non-stochastic mapping. This allows us to stay in a rather simple framework and to propose a review with concrete nu…
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…
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 uses a simple optogenetic model to compare the timing distortion between a randomly-generated target spike sequence and an externally-stimulated neuron spike sequence. The distor…
We construct a model that predicts the statistical properties of spike trains generated by a sensory neuron. The model describes the combined effects of the neuron's intrinsic properties, the noise in the surrounding, and the external driving stimulus. We show that the spike trains exhibit universal statistical behavior over short times, modulated by a strongly stimulus-dependent behavior over long times. These predictions are confirmed in experiments on H1, a motion-sensitive neuron in the fly visual system.
The brain has the phenomenal ability to reorganize itself by forming new connections among neurons and by pruning others. The so-called neural or brain plasticity facilitates the modification of brain structure and function over different time scales. Plasticity might occur due to external stimuli received from the environment, during recovery from brain injury, or due to modifications within the body and brain itself. In this paper, we study the combined effect of short-term (STP) and spike-timing-dependent plasticities (STDP) on the synaptic strength of excitatory coupled Hodgkin-Huxley neur…
Statistical properties of spike trains measured from a sensory neuron in-vivo are studied experimentally and theoretically. Experiments are performed on an identified neuron in the visual system of the blowfly. It is shown that the spike trains exhibit universal behavior over short time, modulated by a stimulus-dependent envelope over long time. A model of the neuron as a nonlinear oscillator driven by noise and an external stimulus, is suggested to account for these results. The model enables a theoretic distinction of the effects of internal neuronal properties from effects of external stimu…
Memristors have been suggested as neuromorphic computing elements. Spike-time dependent plasticity and the Hodgkin-Huxley model of the neuron have both been modelled effectively by memristor theory. The d.c. response of the memristor is a current spike. Based on these three facts we suggest that memristors are well-placed to interface directly with neurons. In this paper we show that connecting a spiking memristor network to spiking neuronal cells causes a change in the memristor network dynamics by: removing the memristor spikes, which we show is due to the effects of connection to aqueous me…
