Recurrent neural networks are powerful tools for understanding and modeling computation and representation by populations of neurons. Continuous-variable or "rate" model networks have been analyzed and applied extensively for these purposes. However, neurons fire action potentials, and the discrete nature of spiking is an important feature of neural circuit dynamics. Despite significant advances, training recurrently connected spiking neural networks remains a challenge. We present a procedure for training recurrently connected spiking networks to generate dynamical patterns autonomously, to produce complex temporal outputs based on integrating network input, and to model physiological data. Our procedure makes use of a continuous-variable network to identify targets for training the inputs to the spiking model neurons. Surprisingly, we are able to construct spiking networks that duplicate tasks performed by continuous-variable networks with only a relatively minor expansion in the number of neurons. Our approach provides a novel view of the significance and appropriate use of "firing rate" models, and it is a useful approach for building model spiking networks that can be used to address important questions about representation and computation in neural systems.
Fast-spiking (FS) interneurons in the brain are self-innervated by powerful inhibitory GABAergic autaptic connections. By computational modelling, we investigate how autaptic inhibition regulates the firing response of such interneurons. Our results indicate that autaptic inhibition both boosts the current threshold for action potential generation as well as modulates the input-output gain of FS interneurons. The autaptic transmission delay is identified as a key parameter that controls the firing patterns and determines multistability regions of FS interneurons. Furthermore, we observe that n…
Heterogeneity of neural attributes has recently gained a lot of attention and is increasing recognized as a crucial feature in neural processing. Despite its importance, this physiological feature has traditionally been neglected in theoretical studies of cortical neural networks. Thus, there is still a lot unknown about the consequences of cellular and circuit heterogeneity in spiking neural networks. In particular, combining network or synaptic heterogeneity and intrinsic heterogeneity has yet to be considered systematically despite the fact that both are known to exist and likely have signi…
The importance of self-feedback autaptic transmission in modulating spike-time irregularity is still poorly understood. By using a biophysical model that incorporates autaptic coupling, we here show that self-innervation of neurons participates in the modulation of irregular neuronal firing, primarily by regulating the occurrence frequency of burst firing. In particular, we find that both excitatory and electrical autapses increase the occurrence of burst firing, thus reducing neuronal firing regularity. In contrast, inhibitory autapses suppress burst firing and therefore tend to improve the r…
We consider a wide class of spiking neuron models, defined by rather general set of conditions typical for basic models like leaky integrate and fire, or binding neuron model. A neuron is fed with a point renewal process. A relation between the three probability density functions (pdf): (i) pdf of input interspike intervals, (ii) pdf of output interspike intervals of a neuron with instantaneous feedback and (iii) pdf for that same neuron without feedback is derived. This allows to calculate any of the three pdfs provided the another two are given. Similar relation between corresponding means a…
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.
Strong inhibitory input to neurons, which occurs in balanced states of neural networks, increases synaptic current fluctuations. This has led to the assumption that inhibition contributes to the high spike-firing irregularity observed in vivo. We used single compartment neuronal models with time-correlated (due to synaptic filtering) and state-dependent (due to reversal potentials) input to demonstrate that inhibitory input acts to decrease membrane potential fluctuations, a result that cannot be achieved with simplified neural input models. To clarify the effects on spike-firing regularity, w…
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.
Periodic and nonperiodic components of electrophysiological signals are modelled in terms of syncronized sequences of closed loops of firing neurons correlated in Markov chains. Single closed loops of firing neurons reproduce fundamental and harmonic components, appearing as lines in the power spectra at frequencies ranging about from $0.5 Hz$ to $ 100 Hz$. Further interesting features of the brainwave signals emerge by considering multiple syncronized sequences of closed loops. In particular, we show that the fluctuations of the number of syncronized loops leads to the onset of broadband powe…
A central question in neuroscience is to understand how noisy firing patterns are used to transmit information. Because neural spiking is noisy, spiking patterns are often quantified via pairwise correlations, or the probability that two cells will spike coincidentally, above and beyond their baseline firing rate. One observation frequently made in experiments, is that correlations can increase systematically with firing rate. Theoretical studies have determined that stimulus-dependent correlations that increase with firing rate can have beneficial effects on information coding; however, we st…
The dominant modeling framework for understanding cortical computations are heuristic firing rate models. Despite their success, these models fall short to capture spike synchronization effects, to link to biophysical parameters and to describe finite-size fluctuations. In this opinion article, we propose that the refractory density method (RDM), also known as age-structured population dynamics or quasi-renewal theory, yields a powerful theoretical framework to build rate-based models for mesoscopic neural populations from realistic neuron dynamics at the microscopic level. We review recent ad…
Effects of distractions such as noises and parameter heterogeneity have been studied on the firing activity of ensemble neurons, each of which is described by the extended Morris-Lecar model showing the graded persisting firings with the aid of an included ${\rm Ca}^{2+}$-dependent cation current. Although the sustained activity of {\it single} neurons is rather robust in a sense that the activity is realized even in the presence of the distractions, the graded frequency of sustained firings is vulnerable to them. It has been shown, however, that the graded persisting activity of {\it ensemble…
We examine the effects of stochastic input currents on the firing behavior of two excitable neurons coupled with fast excitatory synapses. In such cells (models), typified by the quadratic integrate and fire model, mutual synaptic coupling can cause sustained firing or oscillatory behavior which is necessarily antiphase. Additive Gaussian white noise can transiently terminate the oscillations, hence destroying the stable limit cycle. Further application of the noise may return the system to spiking activity. In a particular noise range, the transition times between the oscillating and the rest…
Maintaining the ability to fire sparsely is crucial for information encoding in neural networks. Additionally, spiking homeostasis is vital for spiking neural networks with changing numbers of weights and neurons. We discuss a range of network stabilisation approaches, inspired by homeostatic synaptic plasticity mechanisms reported in the brain. These include weight scaling, and weight change as a function of the network's spiking activity. We tested normalisation of the sum of weights for all neurons, and by neuron type. We examined how this approach affects firing rate and performance on clu…
Starting from a spectral expansion of the Fokker-Plank equation for the membrane potential density in a network of spiking neurons, a low-dimensional dynamics of the collective firing rate is derived. As a result a $n$-order ordinary differential equation for the network activity can be worked out by taking into account the slowest $n$ modes of the expansion. The resulting low-dimensional dynamics naturally takes into account the strength of the synaptic couplings under the hypothesis of a not too fast changing membrane potential density. By considering only the two slowest modes, the firing r…
When attention is directed into the receptive field of a V4 neuron, its contrast response curve is shifted to lower contrast values (Reynolds et al, 2000, Neuron 26:703). Attention also increases the coherence between neurons responding to the same stimulus (Fries et al, 2001, Science 291:1560). We studied how the firing rate and synchrony of a densely interconnected cortical network varied with contrast and how they were modulated by attention. We found that an increased driving current to the excitatory neurons increased the overall firing rate of the network, whereas variation of the drivin…
Correlation-based Hebbian plasticity is thought to shape neuronal connectivity during development and learning, whereas homeostatic plasticity would stabilize network activity. Here we investigate another, new aspect of this dichotomy: Can Hebbian associative properties also emerge as a network effect from a plasticity rule based on homeostatic principles on the neuronal level? To address this question, we simulated a recurrent network of leaky integrate-and-fire neurons, in which excitatory connections are subject to a structural plasticity rule based on firing rate homeostasis. We show that …
In this work we consider a periodically forced generic integrate-and-fire model with a unique attracting equilibrium in the subthreshold dynamics and study the dependence of the firing-rate on the frequency of the drive. In an earlier study we have obtained rigorous results on the bifurcation structure in such systems, with emphasis on the relation between the firing-rate and the rotation number of the existing periodic orbits. In this work we study how these bifurcation structures behave upon variation of the frequency of the input. This allows us to show that the dependence of the firing-rat…
Neuronal models based on the Hodgkin-Huxley equation form a fundamental framework in the field of computational neuroscience. While the neuronal state is often modeled deterministically, experimental recordings show stochastic fluctuations, presumably driven by molecular noise from the underlying microphysical conditions. In turn, the firing of individual neurons gives rise to an electric field n extracellular space, also thought to affect the firing pattern of nearby neurons. We develop a multiscale model which combines a stochastic ion channel gating process taking place on the neuronal memb…
Definition. The phase-of-firing code is a neural coding scheme whereby neurons encode information using the time at which they fire spikes within a cycle of the ongoing oscillatory pattern of network activity. This coding scheme may allow neurons to use their temporal pattern of spikes to encode information that is not encoded in their firing rate.
Neural stimulation represents a powerful technique for neural disorder treatment. This paper deals with optimization of coil design parameters to be used during stimulation for modulation of neuronal firing to achieve pain relief. Pain mechanism is briefly introduced and a hierarchical stimulation model from coil stimulation to neuronal firing is proposed. Electromagnetic field distribution for circular, figure of 8 and Magnetic resonance coupling figure of 8 coils are analyzed with respect to the variation of stimulation parameters such as distance between coils, stimulation frequency, number…
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 …
The coding properties of cells with different types of receptive fields have been studied for decades. ON-type neurons fire in response to positive fluctuations of the time-dependent stimulus, whereas OFF cells are driven by negative stimulus segments. Biphasic cells, in turn, are selective to up/down or down/up stimulus upstrokes. In this paper, we explore the way in which different receptive fields affect the firing statistics of Poisson neuron models, when driven with slow stimuli. We find analytical expressions for the time-dependent peri-stimulus time histogram and the inter-spike interva…
Two recent experimental observations pose a challenge to many cortical models. First, the activity in the auditory cortex is sparse, and firing rates can be described by a lognormal distribution. Second, the distribution of non-zero synaptic strengths between nearby cortical neurons can also be described by a lognormal distribution. Here we use a simple model of cortical activity to reconcile these observations. The model makes the experimentally testable prediction that synaptic efficacies onto a given cortical neuron are statistically correlated, i.e. it predicts that some neurons receive ma…
Neuronal voltage dynamics of regularly firing neurons typically has one stable attractor: either a fixed point (like in the subthreshold regime) or a limit cycle that defines the tonic firing of action potentials (in the suprathreshold regime). In two of the three spike onset bifurcation sequences that are known to give rise to all-or-none type action potentials, however, the resting-state fixpoint and limit cycle spiking can coexist in an intermediate regime, resulting in bistable dynamics. Here, noise can induce switches between the attractors, i.e., between rest and spiking, and thus increa…
The effect of environmental temperature on neuronal spiking behaviors is investigated by numerically simulating the temperature dependence of spiking threshold of the Hodgkin-Huxley neuron subject to synaptic stimulus. We find that the spiking threshold exhibits a global minimum in a "comfortable temperature" range where spike initiation needs weakest synaptic strength, indicating the occurrence of optimal use of synaptic transmission in neural system. We further explore the biophysical origin of this phenomenon in ion channel gating kinetics and also discuss its possible biological relevance …
Repeated occurrences of serial firing sequences of a group of neurons with fixed time delays between neurons are observed in many experiments involving simultaneous recordings from multiple neurons. Such temporal patterns are potentially indicative of underlying microcircuits and it is important to know when a repeatedly occurring pattern is statistically significant. These sequences are typically identified through correlation counts, such as in the two-tape algorithm of Abeles and Gerstein. In this paper we present a method for deciding on the significance of such correlations by characteriz…
We present a simple Markov model of spiking neural dynamics that can be analytically solved to characterize the stochastic dynamics of a finite-size spiking neural network. We give closed-form estimates for the equilibrium distribution, mean rate, variance and autocorrelation function of the network activity. The model is applicable to any network where the probability of firing of a neuron in the network only depends on the number of neurons that fired in a previous temporal epoch. Networks with statistically homogeneous connectivity and membrane and synaptic time constants that are not exces…
A common approach to interpreting spiking activity is based on identifying the firing fields---regions in physical or configuration spaces that elicit responses of neurons. Common examples include hippocampal place cells that fire at preferred locations in the navigated environment, head direction cells that fire at preferred orientations of the animal's head, view cells that respond to preferred spots in the visual field, etc. In all these cases, firing fields were discovered empirically, by trial and error. We argue that the existence and a number of properties of the firing fields can be es…
Humans and other animals behave as if we perform fast Bayesian inference underlying decisions and movement control given uncertain sense data. Here we show that a biophysically realistic model of the subthreshold membrane potential of a single neuron can exactly compute the numerator in Bayes rule for inferring the Poisson parameter of a sensory spike train. A simple network of spiking neurons can construct and represent the Bayesian posterior density of a parameter of an external cause that affects the Poisson parameter, accurately and in real time.
We present a complete mean field theory for a balanced state of a simple model of an orientation hypercolumn. The theory is complemented by a description of a numerical procedure for solving the mean-field equations quantitatively. With our treatment, we can determine self-consistently both the firing rates and the firing correlations, without being restricted to specific neuron models. Here, we solve the analytically derived mean-field equations numerically for integrate-and-fire neurons. Several known key properties of orientation selective cortical neurons emerge naturally from the descript…
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…
Spiking neural networks (SNNs) has attracted much attention due to its great potential of modeling time-dependent signals. The firing rate of spiking neurons is decided by control rate which is fixed manually in advance, and thus, whether the firing rate is adequate for modeling actual time series relies on fortune. Though it is demanded to have an adaptive control rate, it is a non-trivial task because the control rate and the connection weights learned during the training process are usually entangled. In this paper, we show that the firing rate is related to the eigenvalue of the spike gene…
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…
Mounting evidence in recent years suggests that astrocytes, a sub-type of glial cells, not only serve metabolic and structural support for neurons and synapses but also play critical roles in regulation of proper functioning of the nervous system. In this work, we investigate the effect of astrocyte on the spontaneous firing activity of a neuron through a combined model which includes a neuron-astrocyte pair. First, we show that an astrocyte may provide a kind of multistability in neuron dynamics by inducing different firing modes such as random and bursty spiking. Then, we identify the underl…
We study the effect of intrinsic heterogeneity on the activity of a population of leaky integrate-and-fire neurons. By rescaling the dynamical equation, we derive mathematical relations between multiple neuronal parameters and a fluctuating input noise. To this end, common input to heterogeneous neurons is conceived as an identical noise with neuron-specific mean and variance. As a consequence, the neuronal output rates can differ considerably, and their relative spike timing becomes desynchronized. This theory can quantitatively explain some recent experimental findings.
The role of synchronous firing in sensory coding and cognition remains controversial. While studies, focusing on its mechanistic consequences in attentional tasks, suggest that synchronization dynamically boosts sensory processing, others failed to find significant synchronization levels in such tasks. We attempt to understand both lines of evidence within a coherent theoretical framework. We conceptualize synchronization as an independent control parameter to study how the postsynaptic neuron transmits the average firing activity of a presynaptic population, in the presence of synchronization…
Finite-sized populations of spiking elements are fundamental to brain function, but also used in many areas of physics. Here we present a theory of the dynamics of finite-sized populations of spiking units, based on a quasi-renewal description of neurons with adaptation. We derive an integral equation with colored noise that governs the stochastic dynamics of the population activity in response to time-dependent stimulation and calculate the spectral density in the asynchronous state. We show that systems of coupled populations with adaptation can generate a frequency band in which sensory inf…
Autapses are synapses that connect a neuron to itself in the nervous system. Previously, both experimental and theoretical studies have demonstrated that autaptic connections in the nervous system have a significant physiological function. Autapses in nature provide self-delayed feedback, thus introducing an additional time scale to neuronal activities and causing many dynamic behaviors in neurons. Recently, theoretical studies have revealed that an autapse provides a control option for adjusting the response of a neuron: e.g., an autaptic connection can cause the electrical activities of the …
We investigated the influence of efficacy of synaptic interaction on firing synchronization in excitatory neuronal networks. We found spike death phenomena, namely, the state of neurons transits from limit cycle to fixed point or transient state. The phenomena occur under the perturbation of excitatory synaptic interaction that has a high efficacy. We showed that the decrease of synaptic current results in spike death through depressing the feedback of sodium ionic current. In the networks with spike death property the degree of synchronization is lower and unsensitive to the heterogeneity of …
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…
Heterogeneity of firing rate statistics is known to have severe consequences on neural coding. Recent experimental recordings in weakly electric fish indicate that the distribution-width of superficial pyramidal cell firing rates (trial- and time-averaged) in the electrosensory lateral line lobe (ELL) depends on the stimulus, and also that network inputs can mediate changes in the firing rate distribution across the population. We previously developed theoretical methods to understand how two attributes (synaptic and intrinsic heterogeneity) interact and alter the firing rate distribution in a…
Primary vestibular neurons are categorized as either regularly or irregularly firing afferents that use rate and temporal sensory encoding strategies, respectively. While many factors influence firing in these neurons, recent work in mammalian vestibular afferents has demonstrated a rich diversity in ion channels that drive spiking regularity. Here, I review key ionic currents studied in vitro and demonstrate how they may enable sensory encoding strategies demonstrated in vivo.
Even a single neuron may be able to produce significant lognormal features in its firing statistics due to noise in the charging ion current. A mathematical scheme introduced in advanced nanotechnology is relevant for the analysis of this mechanism in the simplest case, the integrate-and-fire model with white noise in the charging ion current.
Networks of model neurons with balanced recurrent excitation and inhibition produce irregular and asynchronous spiking activity. We extend the analysis of balanced networks to include the known dependence of connection probability on the spatial separation between neurons. In the continuum limit we derive that stable, balanced firing rate solutions require that the spatial spread of external inputs be broader than that of recurrent excitation, which in turn must be broader than or equal to that of recurrent inhibition. For finite size networks we investigate the pattern forming dynamics arisin…
A long tradition in theoretical neuroscience casts sensory processing in the brain as the process of inferring the maximally consistent interpretations of imperfect sensory input. Recently it has been shown that Gamma-band inhibition can enable neural attractor networks to approximately carry out such a sampling mechanism. In this paper we propose a novel neural network model based on irregular gating inhibition, show analytically how it implements a Monte-Carlo Markov Chain (MCMC) sampler, and describe how it can be used to model networks of both neural attractors as well as of single spiking…
The response of a neural cell to an external stimulus can follow one of the two patterns: Nonresonant neurons monotonously relax to the resting state after excitation while resonant ones show subthreshold oscillations. We investigate how do these subthreshold properties of neurons affect their suprathreshold response. Vice versa we ask: Can we distinguish between both types of neuronal dynamics using suprathreshold spike trains? The dynamics of neurons is given by stochastic FitzHugh-Nagumo and Morris-Lecar models with either having a focus or a node as the stable fixpoint. We determine numeri…
Many central neurons, and in particular certain brainstem aminergic neurons exhibit spontaneous and fairly regular spiking with frequencies of order a few Hz. A large number of ion channel types contribute to such spiking so that accurate modeling of spike generation leads to the requirement of solving very large systems of differential equations, ordinary in the first instance. Since analysis of spiking behavior when many synaptic inputs are active adds further to the number of components, it is useful to have simplified mathematical models of spiking in such neurons so that, for example, sto…
Chemical and electrical synapses shape the dynamics of neuronal networks. Numerous theoretical studies have investigated how each of these types of synapses contributes to the generation of neuronal oscillations, but their combined effect is less understood. This limitation is further magnified by the impossibility of traditional neuronal mean-field models ---also known as firing rate models, or firing rate equations--- to account for electrical synapses. Here we introduce a novel firing rate model that exactly describes the mean field dynamics of heterogeneous populations of quadratic integra…
Neurons in the intact brain receive a continuous and irregular synaptic bombardment from excitatory and inhibitory pre-synaptic neurons, which determines the firing activity of the stimulated neuron. In order to investigate the influence of inhibitory stimulation on the firing time statistics, we consider Leaky Integrate-and-Fire neurons subject to inhibitory instantaneous post-synaptic potentials. In particular, we report exact results for the firing rate, the coefficient of variation and the spike train spectrum for various synaptic weight distributions. Our results are not limited to stimul…
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…
A population of firing neurons is expected to carry information not only by mean firing rate but also by fluctuation and synchrony among neurons. In order to examine this possibility, we have studied responses of neuronal ensembles to three kinds of inputs: mean-, fluctuation- and synchrony-driven inputs. The generalized rate-code model including additive and multiplicative noise (H. Hasegawa, Phys. Rev. E {\bf 75} (2007) 051904) has been studied by direct simulations (DSs) and the augmented moment method (AMM) in which equations of motion for mean firing rate, fluctuation and synchrony are de…
