We study a network model of two conductance-based pacemaker neurons of differing natural frequency, coupled with either mutual excitation or inhibition, and receiving shared random inhibitory synaptic input. The networks may phase-lock spike-to-spike for strong mutual coupling. But the shared input can desynchronize the locked spike-pairs by selectively eliminating the lagging spike or modulating its timing with respect to the leading spike depending on their separation time window. Such loss of synchrony is also found in a large network of sparsely coupled heterogeneous spiking neurons receiving shared input.
The behaviour of neurons under the influence of periodic external input has been modelled very successfully by circle maps. The aim of this note is to extend certain aspects of this analysis to a much more general class of forcing processes. We apply results on the fibred rotation number of randomly forced circle maps to show the uniqueness of the asymptotic firing frequency of ergodically forced pacemaker neurons. The details of the analysis are carried out for the forced leaky integrate-and-fire model, but the results should also remain valid for a large class of further models.
Serotonergic, noradrenergic and dopaminergic brainstem (including midbrain) neurons, often exhibit spontaneous and fairly regular spiking with frequencies of order a few Hz, though dopaminergic and noradrenergic neurons only exhibit such pacemaker-type activity in vitro or in vivo under special conditions. A large number of ion channel types contribute to such spiking so that detailed modeling of spike generation leads to the requirement of solving very large systems of differential equations. It is useful to have simplified mathematical models of spiking in such neurons so that, for example, …
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…
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…
We study the effects of periodic subthreshold pacemaker activity and time-delayed coupling on stochastic resonance over scale-free neuronal networks. As the two extreme options, we introduce the pacemaker respectively to the neuron with the highest degree and to one of the neurons with the lowest degree within the network, but we also consider the case when all neurons are exposed to the periodic forcing. In the absence of delay, we show that an intermediate intensity of noise is able to optimally assist the pacemaker in imposing its rhythm on the whole ensemble, irrespective to its placing, t…
There is consensus in the current literature that stable states of asynchronous irregular spiking activity require (i) large networks of 10 000 or more neurons and (ii) external background activity or pacemaker neurons. Yet already in 1963, Griffith showed that networks of simple threshold elements can be persistently active at intermediate rates. Here, we extend Griffith's work and demonstrate that sparse networks of integrate-and-fire neurons assume stable states of self-sustained asynchronous and irregular firing without external input or pacemaker neurons. These states can be robustly indu…
Spike-timing dependent plasticity (STDP) is an organizing principle of biological neural networks. While synchronous firing of neurons is considered to be an important functional block in the brain, how STDP shapes neural networks possibly toward synchrony is not entirely clear. We examine relations between STDP and synchronous firing in spontaneously firing neural populations. Using coupled heterogeneous phase oscillators placed on initial networks, we show numerically that STDP prunes some synapses and promotes formation of a feedforward network. Eventually a pacemaker, which is the neuron w…
Spike-timing-dependent plasticity (STDP) with asymmetric learning windows is commonly found in the brain and useful for a variety of spike-based computations such as input filtering and associative memory. A natural consequence of STDP is establishment of causality in the sense that a neuron learns to fire with a lag after specific presynaptic neurons have fired. The effect of STDP on synchrony is elusive because spike synchrony implies unitary spike events of different neurons rather than a causal delayed relationship between neurons. We explore how synchrony can be facilitated by STDP in osc…
Networks of excitatory and inhibitory neurons display asynchronous irregular (AI) states, where the activities of the two populations are balanced. At the single cell level, it was shown that neurons subject to balanced and noisy synaptic inputs can display enhanced responsiveness. We show here that this enhanced responsiveness is also present at the network level, but only when single neurons are in a conductance state and fluctuation regime consistent with experimental measurements. In such states, the entire population of neurons is globally influenced by the external input. We suggest that…
Although recent neurophysiological experiments suggest that synchronous neural activity is involved in some perceptual and cognitive processes, the functional role of such coherent neuronal behavior is not well understood. As a first step in clarifying this role, we investigate how the temporal coherence of certain neuronal activity affects the activity pattern in a neural network. Using a simple network of leaky integrate-and-fire neurons, we study the effects of synchronized incoming spikes on the functioning of two mechanisms typically used in model neural systems, winner-take-all competiti…
We investigate the possibility that narrowband oscillations may emerge from completely asynchronous, independent neural firing. We find that a population of asynchronous neurons may produce narrowband oscillations if each neuron fires quasi-periodically, and we deduce bounds on the degree of variability in neural spike-timing which will permit the emergence of such oscillations. These results suggest a novel mechanism of neural rhythmogenesis, and they help to explain recent experimental reports of large-amplitude local field potential oscillations in the absence of neural spike-timing synchro…
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…
Serotonergic neurons of the dorsal raphe nuclei, with their extensive innervation of nearly the whole brain have important modulatory effects on many cognitive and physiological processes. They play important roles in clinical depression and other psychiatric disorders. In order to quantify the effects of serotonergic transmission on target cells it is desirable to construct computational models and to this end these it is necessary to have details of the biophysical and spike properties of the serotonergic neurons. Here several basic properties are reviewed with data from several studies sinc…
Asynchronous Bioplausible Neuron for SNN for Event Vision
Department of Computer Science, Kingston University, London, UK（计算机科学系，金斯顿大学，伦敦，英国）；Advanced Research and Innovation Center (ARIC), Khalifa University, Abu Dhabi, UAE（先进研究与创新中心（ARIC），哈利法大学，阿布扎赫德，阿联酋）；Department of Aerospace Engineering, Khalifa University of Science and Technology, Abu Dhabi, UAE（航空航天工程系，哈利法科学技术大学，阿布扎赫德，阿联酋）；Ipsotek, an Eviden Company, London（Ipsotek，一家Eviden公司，伦敦）
The study of balanced networks of excitatory and inhibitory neurons has led to several open questions. On the one hand it is yet unclear whether the asynchronous state observed in the brain is autonomously generated, or if it results from the interplay between external drivings and internal dynamics. It is also not known, which kind of network variabilities will lead to irregular spiking and which to synchronous firing states. Here we show how isolated networks of purely excitatory neurons generically show asynchronous firing whenever a minimal level of structural variability is present togeth…
Spontaneous cortical population activity exhibits a multitude of oscillatory patterns, which often display synchrony during slow-wave sleep or under certain anesthetics and stay asynchronous during quiet wakefulness. The mechanisms behind these cortical states and transitions among them are not completely understood. Here we study spontaneous population activity patterns in random networks of spiking neurons of mixed types modeled by Izhikevich equations. Neurons are coupled by conductance-based synapses subject to synaptic noise. We localize the population activity patterns on the parameter d…
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…
We revisit the dynamics of a prototypical model of balanced activity in networks of spiking neutrons. A detailed investigation of the thermodynamic limit for fixed density of connections (massive coupling) shows that, when inhibition prevails, the asymptotic regime is not asynchronous but rather characterized by a self-sustained irregular, macroscopic (collective) dynamics. So long as the connectivity is massive, this regime is found in many different setups: leaky as well as quadratic integrate-and-fire neurons; large and small coupling strength; weak and strong external currents.
We consider here a single-compartment model of these neurons which is capable of describing many of the known features of spike generation, particularly the slow rhythmic pacemaking activity often observed in these cells in a variety of species. Included in the model are ten kinds of voltage dependent ion channels as well as calcium-dependent potassium current. Calcium dynamics includes buffering and pumping. In sections 3-9, each component is considered in detail and parameters estimated from voltage clamp data where possible. In the next two sections simplified versions of some components ar…
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…
The activity of a sparse network of leaky integrate-and-fire neurons is carefully revisited with reference to a regime of a bona-fide asynchronous dynamics. The study is preceded by a finite-size scaling analysis, carried out to identify a setup where collective synchronization is negligible. The comparison between quenched and annealed networks reveals the emergence of substantial differences when the coupling strength is increased, via a scenario somehow reminiscent of a phase transition. For sufficiently strong synaptic coupling, quenched networks exhibit a highly bursting neural activity, …
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…
We study a network of spiking neurons with heterogeneous excitabilities connected via inhibitory delayed pulses. For globally coupled systems the increase of the inhibitory coupling reduces the number of firing neurons by following a Winner Takes All mechanism. For sufficiently large transmission delay we observe the emergence of collective oscillations in the system beyond a critical coupling value. Heterogeneity promotes neural inactivation and asynchronous dynamics and its effect can be counteracted by considering longer time delays. In sparse networks, inhibition has the counterintuitive e…
Networks in the brain consist of different types of neurons. Here we investigate the influence of neuron diversity on the dynamics, phase space structure and computational capabilities of spiking neural networks. We find that already a single neuron of a different type can qualitatively change the network dynamics and that mixed networks may combine the computational capabilities of ones with a single neuron type. We study inhibitory networks of concave leaky (LIF) and convex "anti-leaky" (XIF) integrate-and-fire neurons that generalize irregularly spiking non-chaotic LIF neuron networks. Endo…
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…
The time elapsed model describes the firing activity of an homogeneous assembly of neurons thanks to the distribution of times elapsed since the last discharge. It gives a mathematical description of the probability density of neurons structured by this time. In an earlier work, based on generalized relative entropy methods, it is proved that for highly or weakly connected networks the model exhibits relaxation to the steady state and for moderately connected networks it is obtained numerical evidence of appearance of self-sustained periodic solutions. Here, we go further and, using the partic…
We investigate a network of integrate-and-fire neurons characterized by a distribution of spiking frequencies. Upon increasing the coupling strength, the model exhibits a transition from an asynchronous regime to a nontrivial collective behavior. At variance with the Kuramoto model, (i) the macroscopic dynamics is irregular even in the thermodynamic limit, and (ii) the microscopic (single-neuron) evolution is linearly stable.
We investigate the performance of sparsely-connected networks of integrate-and-fire neurons for ultra-short term information processing. We exploit the fact that the population activity of networks with balanced excitation and inhibition can switch from an oscillatory firing regime to a state of asynchronous irregular firing or quiescence depending on the rate of external background spikes. We find that in terms of information buffering the network performs best for a moderate, non-zero, amount of noise. Analogous to the phenomenon of stochastic resonance the performance decreases for higher a…
Neurons in the primary visual cortex are more or less selective for the orientation of a light bar used for stimulation. A broad distribution of individual grades of orientation selectivity has in fact been reported in all species. A possible reason for emergence of broad distributions is the recurrent network within which the stimulus is being processed. Here we compute the distribution of orientation selectivity in randomly connected model networks that are equipped with different spatial patterns of connectivity. We show that, for a wide variety of connectivity patterns, a linear theory bas…
The population activity of random networks of excitatory and inhibitory leaky integrate-and-fire (LIF) neurons has been studied extensively. In particular, a state of asynchronous activity with low firing rates and low pairwise correlations emerges in sparsely connected networks. We apply linear response theory to evaluate the influence of detailed network structure on neuron dynamics. It turns out that pairwise correlations induced by direct and indirect network connections can be related to the matrix of direct linear interactions. Furthermore, we study the influence of characteristics of th…
Efficient brain simulation is a scientific grand challenge, a parallel/distributed coding challenge and a source of requirements and suggestions for future computing architectures. Indeed, the human brain includes about 10^15 synapses and 10^11 neurons activated at a mean rate of several Hz. Full brain simulation poses Exascale challenges even if simulated at the highest abstraction level. The WaveScalES experiment in the Human Brain Project (HBP) has the goal of matching experimental measures and simulations of slow waves during deep-sleep and anesthesia and the transition to other brain stat…
Cortical neurons are characterized by irregular firing and a broad distribution of rates. The balanced state model explains these observations with a cancellation of mean excitatory and inhibitory currents, which makes fluctuations drive firing. In networks of neurons with current-based synapses, the balanced state emerges dynamically if coupling is strong, i.e. if the mean number of synapses per neuron $K$ is large and synaptic efficacy is of order $1/\sqrt{K}$. When synapses are conductance-based, current fluctuations are suppressed when coupling is strong, questioning the applicability of t…
To understand possible strategies of temporal spike coding in the central nervous system, we study functional neuromimetic models of visual processing for static images. We will first present the retinal model which was introduced by Van Rullen and Thorpe and which represents the multiscale contrast values of the image using an orthonormal wavelet transform. These analog values activate a set of spiking neurons which each fire once to produce an asynchronous wave of spikes. According to this model, the image may be progressively reconstructed from this spike wave thanks to regularities in the …
Network models are routinely downscaled compared to nature in terms of numbers of nodes or edges because of a lack of computational resources, often without explicit mention of the limitations this entails. While reliable methods have long existed to adjust parameters such that the first-order statistics of network dynamics are conserved, here we show that limitations already arise if also second-order statistics are to be maintained. The temporal structure of pairwise averaged correlations in the activity of recurrent networks is determined by the effective population-level connectivity. We f…
Mean-field systems have been previously derived for networks of coupled, two-dimensional, integrate-and-fire neurons such as the Izhikevich, adapting exponential (AdEx) and quartic integrate and fire (QIF), among others. Unfortunately, the mean-field systems have a degree of frequency error and the networks analyzed often do not include noise when there is adaptation. Here, we derive a one-dimensional partial differential equation (PDE) approximation for the marginal voltage density under a first order moment closure for coupled networks of integrate-and-fire neurons with white noise inputs. T…
We consider an excitatory random network of leaky integrate-and-fire pulse coupled neurons. The neurons are connected as in a directed Erdös-Renyi graph with average connectivity $<k>$ scaling as a power law with the number of neurons in the network. The scaling is controlled by a parameter $γ$, which allows to pass from massively connected to sparse networks and therefore to modify the topology of the system. At a macroscopic level we observe two distinct dynamical phases: an Asynchronous State (AS) corresponding to a desynchronized dynamics of the neurons and a Partial Synchronization (PS) r…
Synaptic, dendritic and single-cell kinetics generate significant time delays that shape the dynamics of large networks of spiking neurons. Previous work has shown that such effective delays can be taken into account with a rate model through the addition of an explicit, fixed delay [Roxin et al. PRL 238103 (2005)]. Here we extend this work to account for arbitrary symmetric patterns of synaptic connectivity and generic nonlinear transfer functions. Specifically, we conduct a weakly nonlinear analysis of the dynamical states arising via primary instabilities of the asynchronous state. In this …
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…
We investigate the modes of oscillation of heterogeneous ring-networks of quadratic integrate-and-fire neurons with non-local, space-dependent coupling. Perturbations of the equilibrium state with a particular wave number produce transient standing waves with a specific frequency, analogous to those in a tense string. In the neuronal network, the equilibrium corresponds to a spatially homogeneous, asynchronous state. Perturbations of this state excite the network's oscillatory modes, which reflect the interplay of episodes of synchronous spiking with the excitatory-inhibitory spatial interacti…
A rigorous bridge between spiking-level and macroscopic quantities is an on-going and well-developed story for asynchronously firing neurons, but focus has shifted to include neural populations exhibiting varying synchronous dynamics. Recent literature has used the Ott--Antonsen ansatz (2008) to great effect, allowing a rigorous derivation of an order parameter for large oscillator populations. The ansatz has been successfully applied using several models including networks of Kuramoto oscillators, theta models, and integrate-and-fire neurons, along with many types of network topologies. In th…
We investigate the efficient transmission and processing of weak, subthreshold signals in a realistic neural medium in the presence of different levels of the underlying noise. Assuming Hebbian weights for maximal synaptic conductances -- that naturally balances the network with excitatory and inhibitory synapses -- and considering short-term synaptic plasticity affecting such conductances, we found different dynamic phases in the system. This includes a memory phase where population of neurons remain synchronized, an oscillatory phase where transitions between different synchronized populatio…
When two stimuli are present in the receptive field of a V4 neuron, the firing rate response is between the weakest and strongest response elicited by each of the stimuli alone (Reynolds et al, 1999, Journal of Neuroscience 19:1736-1753). When attention is directed towards the stimulus eliciting the strongest response (the preferred stimulus), the response to the pair is increased, whereas the response decreases when attention is directed to the other stimulus (the poor stimulus). These experimental results were reproduced in a model of a V4 neuron under the assumption that attention modulates…
Columbia University（哥伦比亚大学）；Edmond and Lily Safra Center for Brain Sciences（埃德蒙和莉莉·萨弗拉脑科学中心）；Racah Institute of Physics（拉卡物理研究所）；The Hebrew University of Jerusalem（耶路撒冷希伯来大学）；Center for Brain Science（脑科学中心）；Harvard University（哈佛大学）
Neurons and networks in the cerebral cortex must operate reliably despite multiple sources of noise. To evaluate the impact of both input and output noise, we determine the robustness of single-neuron stimulus selective responses, as well as the robustness of attractor states of networks of neurons performing memory tasks. We find that robustness to output noise requires synaptic connections to be in a balanced regime in which excitation and inhibition are strong and largely cancel each other. We evaluate the conditions required for this regime to exist and determine the properties of networks…
In a first step towards the comprehension of neural activity, one should focus on the stability of the various dynamical states. Even the characterization of idealized regimes, such as a perfectly periodic spiking activity, reveals unexpected difficulties. In this paper we discuss a general approach to linear stability of pulse-coupled neural networks for generic phase-response curves and post-synaptic response functions. In particular, we present: (i) a mean-field approach developed under the hypothesis of an infinite network and small synaptic conductances; (ii) a "microscopic" approach whic…
We investigate the dynamical role of inhibitory and highly connected nodes (hub) in synchronization and input processing of leaky-integrate-and-fire neural networks with short term synaptic plasticity. We take advantage of a heterogeneous mean-field approximation to encode the role of network structure and we tune the fraction of inhibitory neurons $f_I$ and their connectivity level to investigate the cooperation between hub features and inhibition. We show that, depending on $f_I$, highly connected inhibitory nodes strongly drive the synchronization properties of the overall network through d…
