We analyze the collective dynamics of hierarchically structured networks of densely connected spiking neurons. These networks of sub-networks may represent interactions between cell assemblies or different nuclei in the brain. The dynamical activity pattern that results from these interactions depends on the strength of synaptic coupling between them. Importantly, the overall dynamics of a brain region in the absence of external input, so called ongoing brain activity, has been attributed to the dynamics of such interactions. In our study, two different network scenarios are considered: a system with one inhibitory and two excitatory sub-networks, and a network representation with three inhibitory sub-networks. To study the effect of synaptic strength on the global dynamics of the network, two parameters for relative couplings between these sub-networks are considered. For each case, a co-dimension two bifurcation analysis is performed and the results have been compared to large-scale network simulations. Our analysis shows that Generalized Lotka-Volterra (GLV) equations, well-known in predator-prey studies, yield a meaningful population-level description for the collective behavior of spiking neuronal interaction, which have a hierarchical structure. In particular, we observed a striking equivalence between the bifurcation diagrams of spiking neuronal networks and their corresponding GLV equations. This study gives new insight on the behavior of neuronal assemblies, and can potentially suggest new mechanisms for altering the dynamical patterns of spiking networks based on changing the synaptic strength between some groups of neurons.
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…
Randomly connected networks of excitatory and inhibitory spiking neurons provide a parsimonious model of neural variability, but are notoriously unreliable for performing computations. We show that this difficulty is overcome by incorporating the well-documented dependence of connection probability on distance. Spatially extended spiking networks exhibit symmetry-breaking bifurcations and generate spatiotemporal patterns that can be trained to perform dynamical computations.
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…
Cortical neural circuits display highly irregular spiking in individual neurons but variably sized collective firing, oscillations and critical avalanches at the population level, all of which have functional importance for information processing. Theoretically, the balance of excitation and inhibition inputs is thought to account for spiking irregularity and critical avalanches may originate from an underlying phase transition. However, the theoretical reconciliation of these multilevel dynamic aspects in neural circuits remains an open question. Herein, we study excitation-inhibition (E-I) b…
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 consider a new class of non Markovian processes with a countable number of interacting components, both in discrete and continuous time. Each component is represented by a point process indicating if it has a spike or not at a given time. The system evolves as follows. For each component, the rate (in continuous time) or the probability (in discrete time) of having a spike depends on the entire time evolution of the system since the last spike time of the component. In discrete time this class of systems extends in a non trivial way both Spitzer's interacting particle systems, which are Mar…
We investigate the dynamics of large-scale interacting neural populations, composed of conductance based, spiking model neurons with modifiable synaptic connection strengths, which are possibly also subjected to external noisy currents. The network dynamics is controlled by a set of neural population probability distributions ($\mathrm{PPD}$)which are constructed along the same lines as in the Klimontovich approach to the kinetic theory of plasmas. An exact non-closed, nonlinear, system of integro-partial differential equations is derived for the $\mathrm{PPD}$s. As is customary, a closing pro…
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…
Neural population equations such as neural mass or field models are widely used to study brain activity on a large scale. However, the relation of these models to the properties of single neurons is unclear. Here we derive an equation for several interacting populations at the mesoscopic scale starting from a microscopic model of randomly connected generalized integrate-and-fire neuron models. Each population consists of 50 -- 2000 neurons of the same type but different populations account for different neuron types. The stochastic population equations that we find reveal how spike-history eff…
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…
Rapid action potential generation --- spiking --- and alternating intervals of spiking and quiescence --- bursting --- are two dynamic patterns observed in neuronal activity. In computational models of neuronal systems, the transition from spiking to bursting often exhibits complex bifurcation structure. One type of transition involves the torus canard, which was originally observed in a simple biophysical model of a Purkinje cell. In this article, we expand on that original result by showing that torus canards arise in a broad array of well-known computational neuronal models with three diffe…
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…
The classical Hodgkin--Huxley (HH) model neglects the time-dependence of ion concentrations in spiking dynamics. The dynamics is therefore limited to a time scale of milliseconds, which is determined by the membrane capacitance multiplied by the resistance of the ion channels, and by the gating time constants. We study slow dynamics in an extended HH framework that includes time-dependent ion concentrations, pumps, and buffers. Fluxes across the neuronal membrane change intra- and extracellular ion concentrations, whereby the latter can also change through contact to reservoirs in the surround…
Collective oscillations and their suppression by external stimulation are analyzed in a large-scale neural network consisting of two interacting populations of excitatory and inhibitory quadratic integrate-and-fire neurons. In the limit of an infinite number of neurons, the microscopic model of this network can be reduced to an exact low-dimensional system of mean-field equations. Bifurcation analysis of these equations reveals three different dynamic modes in a free network: a stable resting state, a stable limit cycle, and bistability with a coexisting resting state and a limit cycle. We sho…
This paper models the dynamics of a large set of interacting neurons within the framework of statistical field theory. We use a method initially developed in the context of statistical field theory [44] and later adapted to complex systems in interaction [45][46]. Our model keeps track of individual interacting neurons dynamics but also preserves some of the features and goals of neural field dynamics, such as indexing a large number of neurons by a space variable. Thus, this paper bridges the scale of individual interacting neurons and the macro-scale modelling of neural field theory.
The brain is composed of complex networks of interacting neurons that express considerable heterogeneity in their physiology and spiking characteristics. How does neural heterogeneity affect macroscopic neural dynamics and how does it contribute to neurodynamic functions? In this letter, we address these questions by studying the macroscopic dynamics of networks of heterogeneous Izhikevich neurons. We derive mean-field equations for these networks and examine how heterogeneity in the spiking thresholds of Izhikevich neurons affects the emergent macroscopic dynamics. Our results suggest that th…
Mean-field theory links the physiological properties of individual neurons to the emergent dynamics of neural population activity. These models provide an essential tool for studying brain function at different scales; however, for their application to neural populations on large scale, they need to account for differences between distinct neuron types. The Izhikevich single neuron model can account for a broad range of different neuron types and spiking patterns, thus rendering it an optimal candidate for a mean-field theoretic treatment of brain dynamics in heterogeneous networks. Here, we d…
We review different aspects of the simulation of spiking neural networks. We start by reviewing the different types of simulation strategies and algorithms that are currently implemented. We next review the precision of those simulation strategies, in particular in cases where plasticity depends on the exact timing of the spikes. We overview different simulators and simulation environments presently available (restricted to those freely available, open source and documented). For each simulation tool, its advantages and pitfalls are reviewed, with an aim to allow the reader to identify which s…
Mean field theory is a device to analyze the collective behavior of a dynamical system comprising many interacting particles. The theory allows to reduce the behavior of the system to the properties of a handful of parameters. In neural circuits, these parameters are typically the firing rates of distinct, homogeneous subgroups of neurons. Knowledge of the firing rates under conditions of interest can reveal essential information on both the dynamics of neural circuits and the way they can subserve brain function. The goal of this chapter is to provide an elementary introduction to the mean fi…
We analyse the dynamics of the improved discretised version of the well known Izhikevich neuronmodel under the action of external electromagnetic field. It is found that the three-dimensional IZHmap shows rich dynamics. With the variation of the electromagnetic field, period-doubling routeto chaos in a repeating fashion is observed from the bifurcation diagram. Even the forward andbackward continuation bifurcation diagram which do not completely overlap suggests that there is multistability in the system. The phenomenon of bistability (coexistence of periodic and chaotic attractors) is observe…
Characterizing metastable neural dynamics in finite-size spiking networks remains a daunting challenge. We propose to address this challenge in the recently introduced replica-mean-field (RMF) limit. In this limit, networks are made of infinitely many replicas of the finite network of interest, but with randomized interactions across replicas. Such randomization renders certain excitatory networks fully tractable at the cost of neglecting activity correlations, but with explicit dependence on the finite size of the neural constituents. However, metastable dynamics typically unfold in networks …
Cross-frequency coupling (CFC) is thought to play an important role in communication across distant brain regions. However, neither the mechanism of its generation nor the influence on the underlying spiking dynamics is well understood. Here, we investigate the dynamics of two interacting distant neuronal modules coupled by inter-regional long-range connections. Each neuronal module comprises an excitatory and inhibitory population of quadratic integrate-and-fire neurons connected locally with conductance-based synapses. The two modules are coupled reciprocally with delays that represent the l…
We propose a biologically plausible computational model of working memory (WM) implemented by the spiking neuron network (SNN) interacting with a network of astrocytes. SNN is modelled by the synaptically coupled Izhikevich neurons with a non-specific architecture connection topology. Astrocytes generating calcium signals are connected by local gap junction diffusive couplings and interact with neurons by chemicals diffused in the extracellular space. Calcium elevations occur in response to the increase of concentration of a neurotransmitter released by spiking neurons when a group of them fir…
To understand the collective spiking activity in neuronal populations, it is essential to reveal basic circuit variables responsible for these emergent functional states. Here, I develop a mean field theory for the population coupling recently proposed in the studies of visual cortex of mouse and monkey, relating the individual neuron activity to the population activity, and extend the original form to the second order, relating neuron-pair's activity to the population activity, to explain the high order correlations observed in the neural data. I test the computational framework on the salama…
Short-Term Synaptic Plasticity (STSP) strongly affects the neural dynamics of cortical networks. The Tsodyks and Markram (TM) model for STSP accurately accounts for a wide range of physiological responses at different types of cortical synapses. Here, we report for the first time a route to chaotic behavior via a Shilnikov homoclinic bifurcation that dynamically organises some of the responses in the TM model. In particular, the presence of such a homoclinic bifurcation strongly affects the shape of the trajectories in the phase space and induces highly irregular transient dynamics; indeed, in…
Regime Mapping of Oscillatory States in Balanced Spiking Networks with Multiple Time Scales
Graduate Institute of Networking and Multimedia, National Taiwan University（台湾大学网际多媒体研究所）；Graduate Institute of Vehicle Engineering, National Changhua University of Education（国立彰化师范大学车辆工程研究所）
Low-dimensional descriptions of neural network dynamics are an effective tool for bridging different scales of organization of brain structure and function. Recent advances in deriving mean-field descriptions for networks of coupled oscillators have sparked the development of a new generation of neural mass models. Of notable interest are mean-field descriptions of all-to-all coupled quadratic integrate-and-fire (QIF) neurons, which have already seen numerous extensions and applications. These extensions include different forms of short-term adaptation (STA) considered to play an important rol…
This paper introduces a class of stochastic models of interacting neurons with emergent dynamics similar to those seen in local cortical populations, and compares them to very simple reduced models driven by the same mean excitatory and inhibitory currents. Discrepancies in firing rates between network and reduced models were investigated, and mechanisms leading to these discrepancies were identified. Chief among them is correlations in spiking, or partial synchronization, working in concert with "nonlinearities" in the time evolution of membrane potentials. Additionally, simple random walk mo…
Populations of spiking neuron models have densities of their microscopic variables (e.g., single-cell membrane potentials) whose evolution fully capture the collective dynamics of biological networks, even outside equilibrium. Despite its general applicability, the Fokker-Planck equation governing such evolution is mainly studied within the borders of the linear response theory, although alternative spectral expansion approaches offer some advantages in the study of the out-of-equilibrium dynamics. This is mainly due to the difficulty in computing the state-dependent coefficients of the expand…
Spike-frequency adaptation (SFA) is a fundamental neuronal mechanism taking into account the fatigue due to spike emissions and the consequent reduction of the firing activity. We have studied the effect of this adaptation mechanism on the macroscopic dynamics of excitatory and inhibitory networks of quadratic integrate-and-fire (QIF) neurons coupled via exponentially decaying post-synaptic potentials. In particular, we have studied the population activities by employing an exact mean field reduction, which gives rise to next generation neural mass models. This low-dimensional reduction allows…
We introduce a nonlinear modification of the classical Hawkes process, which allows inhibitory couplings between units without restrictions. The resulting system of interacting point processes provides a useful mathematical model for recurrent networks of spiking neurons with exponential transfer functions. The expected rates of all neurons in the network are approximated by a first-order differential system. We study the stability of the solutions of this equation, and use the new formalism to implement a winner-takes-all network that operates robustly for a wide range of parameters. Finally,…
This paper contains an analysis of a simple neural network that exhibits self-organized criticality. Such criticality follows from the combination of a simple neural network with an excitatory feedback loop that generates bistability, in combination with an anti-Hebbian synapse in its input pathway. Using the methods of statistical field theory, we show how one can formulate the stochastic dynamics of such a network as the action of a path integral, which we then investigate using renormalization group methods. The results indicate that the network exhibits hysteresis in switching back and for…
Scalable Construction of Spiking Neural Networks using up to thousands of GPUs
University of Cagliari（卡利亚里大学）；Istituto Nazionale di Fisica Nucleare (INFN)（意大利国家核物理研究所）；Jülich Research Centre（于利希研究中心）；RWTH Aachen University（亚琛工业大学）
We analyze a large system of heterogeneous quadratic integrate-and-fire (QIF) neurons with time delayed, all-to-all synaptic coupling. The model is exactly reduced to a system of firing rate equations that is exploited to investigate the existence, stability and bifurcations of fully synchronous, partially synchronous, and incoherent states. In conjunction with this analysis we perform extensive numerical simulations of the original network of QIF neurons, and determine the relation between the macroscopic and microscopic states for partially synchronous states. The results are summarized in t…
We describe a transition from bursting to rapid spiking in a reduced mathematical model of a cerebellar Purkinje cell. We perform a slow-fast analysis of the system and find that -- after a saddle node bifurcation of limit cycles -- the full model dynamics follow temporarily a repelling branch of limit cycles. We propose that the system exhibits a dynamical phenomenon new to realistic, biophysical applications: torus canards.
