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 characterizing the influence of one neuron on another in terms of conditional probabilities and specifying our null hypothesis in terms of a bound on the conditional probabilities. This method of testing significance of correlation counts is more general than the currently available methods since under our null hypothesis we do not assume that the spiking processes of different neurons are independent. The structure of our null hypothesis also allows us to rank order the detected patterns in terms of the strength of interaction among the neurons constituting the pattern. We demonstrate our method of assessing significance on simulated spike trains involving inhomogeneous Poisson processes with strong interactions, where the correlation counts are obtained using the two-tape algorithm.
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…
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…
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 timing-based neural codes, neurons have to emit action potentials at precise moments in time. We use a supervised learning paradigm to derive a synaptic update rule that optimizes via gradient ascent the likelihood of postsynaptic firing at one or several desired firing times. We find that the optimal strategy of up- and downregulating synaptic efficacies can be described by a two-phase learning window similar to that of Spike-Timing Dependent Plasticity (STDP). If the presynaptic spike arrives before the desired postsynaptic spike timing, our optimal learning rule predicts that the synapse…
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…
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.
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…
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 …
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…
Two-photon imaging of calcium indicators allows simultaneous recording of responses of hundreds of neurons over hours and even days, but provides a relatively indirect measure of their spiking activity. Existing deconvolution algorithms attempt to recover spikes from observed imaging data, which are then commonly subjected to the same analyses that are applied to electrophysiologically recorded spikes (e.g., estimation of average firing rates, or tuning curves). Here we show, however, that in the presence of noise this approach is often heavily biased. We propose an alternative analysis that a…
Realizations of low firing rates in neural networks usually require globally balanced distributions among excitatory and inhibitory links, while feasibility of temporal coding is limited by neuronal millisecond precision. We show that cooperation, governing global network features, emerges through nodal properties, as opposed to link distributions. Using in vitro and in vivo experiments we demonstrate microsecond precision of neuronal response timings under low stimulation frequencies, whereas moderate frequencies result in a chaotic neuronal phase characterized by degraded precision. Above a …
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…
In this correspondence information theoretical tools are used to investigate the statistical properties of modeled cochlear nucleus globular bushy cell spike trains. The firing patterns are obtained from a simulation software that generates sample spike trains from any auditory input. Here we analyze for the first time the responses of globular bushy cells to voiced and unvoiced speech sounds. Classical entropy estimates, such as the direct method, are improved upon by considering a time-varying and time-dependent entropy estimate. With this method we investigated the relationship between the …
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…
Neuroscientists formulate very different hypotheses about the nature of neural code. At one extreme, it has been argued that neurons encode information in relatively slow changes of individual spikes arriving "rates codes" and the irregularity in the spike trains reflects noise in the system, while in the other extreme this irregularity is the temporal codes thus the precise timing of every spike carries additional information about the input. It is known that in the estimation of Shannon information the patterns and temporal structures are taken into account, while the rate code is determined…
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…
We investigate the stimulus-dependent tuning properties of a noisy ionic conductance model for intrinsic subthreshold oscillations in membrane potential and associated spike generation. On depolarization by an applied current, the model exhibits subthreshold oscillatory activity with occasional spike generation when oscillations reach the spike threshold. We consider how the amount of applied current, the noise intensity, variation of maximum conductance values and scaling to different temperature ranges alter the responses of the model with respect to voltage traces, interspike intervals and …
In this paper, we examine the effects of correlated Gaussian noise on a two-dimensional neuronal network that is locally modeled by the Rulkov map. More precisely, we study the effects of the noise correlation on the variations of the mean firing rate and the correlations among neurons versus the noise intensity. Via numerical simulations, we show that the mean firing rate can always be optimized at an intermediate noise intensity, irrespective of the noise correlation. On the other hand, variations of the population coherence with respect to the noise intensity are strongly influenced by the …
Cell ensembles, originally proposed by Donald Hebb in 1949, are subsets of synchronously firing neurons and proposed to explain basic firing behavior in the brain. Despite having been studied for many years no conclusive evidence has been presented yet for their existence and involvement in information processing such that their identification is still a topic of modern research, especially since simultaneous recordings of large neuronal population have become possible in the past three decades. These large recordings pose a challenge for methods allowing to identify individual neurons forming…
A classical view of neural coding relies on temporal firing synchrony among functional groups of neurons; however the underlying mechanism remains an enigma. Here we experimentally demonstrate a mechanism where time-lags among neuronal spiking leap from several tens of milliseconds to nearly zero-lag synchrony. It also allows sudden leaps out of synchrony, hence forming short epochs of synchrony. Our results are based on an experimental procedure where conditioned stimulations were enforced on circuits of neurons embedded within a large-scale network of cortical cells in vitro and are corrobor…
A good understanding of how neurons use electrical pulses (i.e, spikes) to encode the signal information remains elusive. Analyzing spike sequences generated by individual neurons and by two coupled neurons (using the stochastic FitzHugh-Nagumo model), recent theoretical studies have found that the relative timing of the spikes can encode the signal information. Using a symbolic method to analyze the spike sequence, preferred and infrequent spike patterns were detected, whose probabilities vary with both, the amplitude and the frequency of the signal. To investigate if this encoding mechanism …
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…
In view of ever-changing conditions both in the external world and in intrinsic brain states, maintaining the robustness of computations poses a challenge, adequate solutions to which we are only beginning to understand. At the level of cell-intrinsic properties, biophysical models of neurons permit one to identify relevant physiological substrates that can serve as regulators of neuronal excitability and to test how feedback loops can stabilize crucial variables such as long-term calcium levels and firing rates. Mathematical theory has also revealed a rich set of complementary computational p…
Our mysterious brain is believed to operate near a non-equilibrium point and generate critical self-organized avalanches in neuronal activity. Recent experimental evidence has revealed significant heterogeneity in both synaptic input and output connectivity, but whether the structural heterogeneity participates in the regulation of neuronal avalanches remains poorly understood. By computational modelling, we predict that different types of structural heterogeneity contribute distinct effects on avalanche neurodynamics. In particular, neuronal avalanches can be triggered at an intermediate leve…
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.
The representation of the natural-density, heterogeneous connectivity of neuronal network models at relevant spatial scales remains a challenge for Computational Neuroscience and Neuromorphic Computing. In particular, the memory demands imposed by the vast number of synapses in brain-scale network simulations constitutes a major obstacle. Limiting the number resolution of synaptic weights appears to be a natural strategy to reduce memory and compute load. In this study, we investigate the effects of a limited synaptic-weight resolution on the dynamics of recurrent spiking neuronal networks res…
The synaptic connectivity of cortex is plastic, with experience shaping the ongoing interactions between neurons. Theoretical studies of spike timing-dependent plasticity (STDP) have focused on either just pairs of neurons or large-scale simulations where analytic insight is lacking. A simple account for how fast spike time correlations affect both micro- and macroscopic network structure remains lacking. We develop a low-dimensional mean field theory showing how STDP gives rise to strongly coupled assemblies of neurons with shared stimulus preferences, with the connectivity actively reinforce…
The hippocampal system contains neural populations that encode an animal's position and velocity as it navigates through space. Here, we show that such populations can embed two codes within their spike trains: a firing rate code (R) conveyed by within-cell spike intervals, and a co-firing rate code (R') conveyed by between-cell spike intervals. These two codes behave as conjugates of one another, obeying an analog of the uncertainty principle from physics: information conveyed in R comes at the expense of information in R', and vice versa. An exception to this trade-off occurs when spike trai…
Recent studies of cortical neurons driven by fluctuating currents revealed cutoff frequencies for action potential encoding of several hundred Hz. Theoretical studies of biophysical neuron models have predicted a much lower cutoff frequency of the order of average firing rate or the inverse membrane time constant. The biophysical origin of the observed high cutoff frequencies is thus not well understood. Here we introduce a neuron model with dynamical action potential generation, in which the linear response can be analytically calculated for uncorrelated synaptic noise. We find that the cutof…
Neurons in the central nervous system communicate with each other with the help of series of Action Potentials, or spike trains. Various studies have shown that neurons encode information in different features of spike trains, such as the fine temporal structure, mean firing rate, synchrony etc. An important step in understanding the encoding of information by neurons, is to obtain a reliable measure of correlation between different spike trains. In this paper, two new binless similarity measures for spike trains are proposed. The performance of the new measures are compared with some existing…
One step in the conventional analysis of extracellularly recorded neuronal data is spike sorting, which separates electrical signal into action potentials from different neurons. Because spike sorting involves human judgment, it can be subjective and time intensive, particularly for large sets of neurons. Here we propose a simple, automated way to construct alternative representations of neuronal activity, called spectral representation (SR). In this approach, neuronal spikes are mapped to a discrete space of spike- waveform features and time. Spectral representation enables us to find single-…
Neuroprosthesis, as one type of precision medicine device, is aiming for manipulating neuronal signals of the brain in a closed-loop fashion, together with receiving stimulus from the environment and controlling some part of our brain/body. In terms of vision, incoming information can be processed by the brain in millisecond interval. The retina computes visual scenes and then sends its output as neuronal spikes to the cortex for further computation. Therefore, the neuronal signal of interest for retinal neuroprosthesis is spike. Closed-loop computation in neuroprosthesis includes two stages: …
Fluorescent nanodiamonds (FND) are carbon-based nanomaterials that can efficiently incorporate optically active photoluminescent centers such as the nitrogen-vacancy complex, thus making them promising candidates as optical biolabels and drug-delivery agents. FNDs exhibit bright fluorescence without photobleaching combined with high uptake rate and low cytotoxicity. Focusing on FNDs interference with neuronal function, here we examined their effect on cultured hippocampal neurons, monitoring the whole network development as well as the electrophysiological properties of single neurons. We obse…
Recurrently coupled networks of inhibitory neurons robustly generate oscillations in the gamma band. Nonetheless, the corresponding Wilson-Cowan type firing rate equation for such an inhibitory population does not generate such oscillations without an explicit time delay. We show that this discrepancy is due to a voltage-dependent spike-synchronization mechanism inherent in networks of spiking neurons which is not captured by standard firing rate equations. Here we investigate an exact low-dimensional description for a network of heterogeneous canonical type-I inhibitory neurons which includes…
We present a modified library-based method for simulating the Hodgkin-Huxley (HH) neuronal networks. By pre-computing a high resolution data library during the interval of an action potential (spike), we can avoid evolving the HH equations during the spike and can use a large time step to raise efficiency. The library method can stably achieve at most 10 times of speedup compared with the regular Runge-Kutta method while capturing most statistical properties of HH neurons like the distribution of spikes which data is widely used in the statistical analysis like transfer entropy and Granger cau…
We study the impact of the decrease in the neural population on the neuronal firing rate. We propose a cellular automaton model from cancerous growth in a brain tissue and the death of neurons due absence of cells that help support to neurons. We use this model to study how the firing rate changes when the neuronal networks is under different external stimuli and the cancerous cells have different proliferation rate. Our work shows that the cancer proliferation decreases the neuronal firing rate.
Neuronal firing activities have attracted a lot of attention since a large population of spatiotemporal patterns in the brain is the basis for adaptive behavior and can also reveal the signs for various neurological disorders including Alzheimer's, schizophrenia, epilepsy and others. Here, we study the dynamics of a simple neuronal network using different sets of settings on a neuromorphic chip. We observed three different types of collective neuronal firing activities, which agree with the clinical data taken from the brain. We constructed a brain phase diagram and showed that within the weak…
Neuronal networks are controlled by a combination of the dynamics of individual neurons and the connectivity of the network that links them together. We study a minimal model of the preBotzinger complex, a small neuronal network that controls the breathing rhythm of mammals through periodic firing bursts. We show that the properties of a such a randomly connected network of identical excitatory neurons are fundamentally different from those of uniformly connected neuronal networks as described by mean-field theory. We show that (i) the connectivity properties of the networks determines the loc…
An essential step toward understanding neural circuits is linking their structure and their dynamics. In general, this relationship can be almost arbitrarily complex. Recent theoretical work has, however, begun to identify some broad principles underlying collective spiking activity in neural circuits. The first is that local features of network connectivity can be surprisingly effective in predicting global statistics of activity across a network. The second is that, for the important case of large networks with excitatory-inhibitory balance, correlated spiking persists or vanishes depending …
In previous articles we have investigated the firing properties of the standard Hodgkin-Huxley (HH) systems of ordinary and partial differential equations in response to input currents composed of a drift (mean) and additive Gaussian white noise. For certain values of the mean current, as the noise amplitude increased from zero, the firing rate exhibited a minimum and this phenomenon was called inverse stochastic resonance (ISR). Here we analyse the underlying transitions from a stable equilibrium point to the limit cycle and vice-versa. Focusing on the case of a mean input current density $μ=…
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…
A common way of studying the relationship between neural activity and behavior is through the analysis of neuronal spike trains that are recorded using one or more electrodes implanted in the brain. Each spike train typically contains spikes generated by multiple neurons. A natural question that arises is "what is the number of neurons $ν$ generating the spike train?"; This article proposes a method-of-moments technique for estimating $ν$. This technique estimates the noise nonparametrically using data from the silent region of the spike train and it applies to isolated spikes with a possibly …
Interval jitter and spike resampling methods are used to analyze the time scale on which temporal correlations occur. They allow the computation of jitter corrected cross correlograms and the performance of an associated statistically robust hypothesis test to decide whether observed correlations at a given time scale are significant. Currently used Monte Carlo methods approximate the probability distribution of coincidences. They require generating $N_{\rm MC}$ simulated spike trains of length $T$ and calculating their correlation with another spike train up to lag $τ_{\max}$. This is computa…
Many types of neurons exhibit spike rate adaptation, mediated by intrinsic slow $\mathrm{K}^+$-currents, which effectively inhibit neuronal responses. How these adaptation currents change the relationship between in-vivo like fluctuating synaptic input, spike rate output and the spike train statistics, however, is not well understood. In this computational study we show that an adaptation current which primarily depends on the subthreshold membrane voltage changes the neuronal input-output relationship (I-O curve) subtractively, thereby increasing the response threshold. A spike-dependent adap…
Statistical similarities between neuronal spike trains could reveal significant information on complex underlying processing. In general, the similarity between synchronous spike trains is somewhat easy to identify. However, the similar patterns also potentially appear in an asynchronous manner. However, existing methods for their identification tend to converge slowly, and cannot be applied to short sequences. In response, we propose Hellinger distance measure based on empirical probabilities, which we show to be as accurate as existing techniques, yet faster to converge for synthetic as well…
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…
Various classes of neurons alternate between high-frequency discharges and silent intervals. This phenomenon is called burst firing. To analyze burst activity in an insect system, grasshopper auditory receptor neurons were recorded in vivo for several distinct stimulus types. The experimental data show that both burst probability and burst characteristics are strongly influenced by temporal modulations of the acoustic stimulus. The tendency to burst, hence, is not only determined by cell-intrinsic processes, but also by their interaction with the stimulus time course. We study this interaction…
How do neuronal populations encode time-dependent stimuli in their population firing rate? To address this question, I consider the quasi-renewal equation and the event-based expansion, two theoretical approximations proposed recently, and test these against peri-stimulus time histograms from L2-3 pyramidal cells in vitro. Parameters are optimized by gradient descent to best match the firing rate output given the current input. The fitting method can estimate single-neuron parameters that are normally obtained either with intracellular recordings or with individual spike trains. I find that qu…
Finite-state Markovian surrogates for long-time neuronal state distributions and firing rates
Courant Institute of Mathematical Sciences, New York University（纽约大学柯朗数学科学研究所）；Center for Bioinformatics, National Laboratory of Protein Engineering and Plant Genetic Engineering, School of Life Sciences, Peking University（北京大学生命科学学院生物信息中心，蛋白质工程及植物基因工程国家重点实验室）；Center for Quantitative Biology, Academy for Advanced Interdisciplinary Studies, Peking University（北京大学前沿交叉学科研究院定量生物学中心）；NYU-ECNU Institute of Brain and Cognitive Science, New York University Shanghai（上海纽约大学脑与认知科学联合研究院）；NYU-ECNU Institute of Mathematical Sciences, New York University Shanghai（上海纽约大学数学科学联合研究院）
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…
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…
