Phase-amplitude coupling (PAC) describes the phenomenon where the power of a high-frequency oscillation evolves with the phase of a low-frequency one. We propose a model that explains the emergence of PAC in two commonly-accepted architectures in the brain, namely, a high-frequency neural oscillation driven by an external low-frequency input and two interacting local oscillations with distinct, locally-generated frequencies. We further propose an interconnection structure for brain regions and demonstrate that low-frequency phase synchrony can integrate high-frequency activities regulated by local PAC and control the direction of information flow across distant regions.
We discuss synchronization in networks of neuronal oscillators which are interconnected via diffusive coupling, i.e. linearly coupled via gap junctions. In particular, we present sufficient conditions for synchronization in these networks using the theory of semi-passive and passive systems. We show that the conductance-based neuronal models of Hodgkin-Huxley, Morris-Lecar, and the popular reduced models of FitzHugh-Nagumo and Hindmarsh-Rose all satisfy a semi-passivity property, i.e. that is the state trajectories of such a model remain oscillatory but bounded provided that the supplied (elec…
Nonlinear oscillators have been used to model neurons that fire periodically in the absence of input. These oscillators, which are called neuronal oscillator, share some common response structures with other biological oscillations. In this study, we analyze the dependence of the global dynamics of an impulse-driven stochastic neuronal oscillator on the relaxation rate to the limit cycle, the strength of the intrinsic noise, and the impulsive input parameters. To do this, we use a Markov operator that both reflects the density evolution of the oscillator and is an extension of the phase transi…
In this paper, we propose a dynamic delayed feedback control approach for desynchronization of chaotic-bursting synchronous activities in an ensemble of globally coupled neuronal oscillators. We demonstrate that the difference signal between an ensemble's mean field and its time delayed state, filtered and fed back to the ensemble, can suppress the self-synchronization in the ensemble. These individual units are decoupled and stabilized at the desired desynchronized states while the stimulation signal reduces to the noise level. The effectiveness of the method is illustrated by examples of two…
Propagation of oscillatory signals through the cortex and coherence is shaped by the connectivity structure of neuronal circuits. This study systematically investigates the network and stimulus properties that shape network responses. The results show how input to a cortical column model of the primary visual cortex excites dynamical modes determined by the laminar pattern. Stimulating the inhibitory neurons in the upper layer reproduces experimentally observed resonances at gamma frequency whose origin can be traced back to two anatomical sub-circuits. We develop this result systematically: I…
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 …
Oscillatory activities are widely observed in specific frequency bands of recorded field potentials in different brain regions, and play critical roles in processing neural information. Understanding the structure of these oscillatory activities is essential for understanding the brain function. So far many details remain elusive about their rhythmic structures and how these oscillations are generated. We show that many oscillatory activities in spontaneous cortical-subcortical networks, such as delta, spindle, gamma, high-gamma and sharp wave ripple bands in different brain regions, are genui…
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 …
Analytical expressions for scaling of brain wave spectra derived from the general nonlinear wave Hamiltonian form show excellent agreement with experimental "neuronal avalanche" data. The theory of the weakly evanescent nonlinear brain wave dynamics reveals the underlying collective processes hidden behind the phenomenological statistical description of the neuronal avalanches and connects together the whole range of brain activity states, from oscillatory wave-like modes, to neuronal avalanches, to incoherent spiking, showing that the neuronal avalanches are just the manifestation of the diff…
The stability of synchronous states is analysed in the context of two populations of inhibitory and excitatory neurons, characterized by different pulse-widths. The problem is reduced to that of determining the eigenvalues of a suitable class of sparse random matrices, randomness being a consequence of the network structure. A detailed analysis, which includes also the study of finite-amplitude perturbations, is performed in the limit of narrow pulses, finding that the stability depends crucially on the relative pulse-width. This has implications for the overall property of the asynchronous (b…
The brain can be understood as a collection of interacting neuronal oscillators, but the extent to which its sustained activity is due to coupling among brain areas is still unclear. Here we study the joint dynamics of two cortical columns described by Jansen-Rit neural mass models, and show that coupling between the columns gives rise to stochastic initiations of sustained collective activity, which can be interpreted as epileptic events. For large enough coupling strengths, termination of these events results mainly from the emergence of synchronization between the columns, and thus is contr…
Chimera states---the coexistence of synchrony and asynchrony in a nonlocally-coupled network of identical oscillators---are often used as a model framework for epileptic seizures. Here, we explore the dynamics of chimera states in a network of modified Hindmarsh-Rose neurons, configured to reflect the graph of the mesoscale mouse connectome. Our model produces superficially epileptiform activity converging on persistent chimera states in a large region of a two-parameter space governing connections (a) between subcortices within a cortex and (b) between cortices. Our findings contribute to a g…
We investigate states of enhanced activity in a biological neuronal network composed of pulse-coupled oscillators. The synaptic couplings between the neurons are dynamic, modeling spike time dependent plasticity. The network exhibits statistical characteristics which recently have been identified in an analysis of epileptic seizures [Osorio et al., Phys. Rev. E 82, 021919(2010)] based on analogies to the onset of earth quakes.
We study an abstracted model of neuronal activity via numerical simulation, and report spatiotemporal pattern formation and critical like dynamics. A population of pulse coupled, discretised, relaxation oscillators is simulated over networks with varying edge density and spatial embedded ness. For intermediate edge density and sufficiently strong spatial embeddedness, we observe a novel spatiotemporal pattern in the field of oscillator phases, visually resembling the surface of a frothing liquid. Increasing the edge density results in critical dynamics, with the distribution of neuronal avalan…
We study synaptically coupled neuronal networks to identify the role of coupling delays in network's synchronized behaviors. We consider a network of excitable, relaxation oscillator neurons where two distinct populations, one excitatory and one inhibitory, are coupled and interact with each other. The excitatory population is uncoupled, while the inhibitory population is tightly coupled. A geometric singular perturbation analysis yields existence and stability conditions for synchronization states under different firing patterns between the two populations, along with formulas for the periods…
While phase oscillators are often used to model neuronal populations, in contrast to the Kuramoto paradigm, strong interactions between brain areas can be associated with loss of synchrony. Using networks of coupled oscillators described by neural mass models, we find that a transition to decoherence at increased coupling strength results from the fundamental nonlinearity, e.g., arising from refractoriness, of the interactions between the nodes. The nonlinearity-driven transition also depends on the connection topology, underlining the role of network structure in shaping brain activity.
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 …
We study numerically and analytically first- and second-order phase transitions in neuronal networks stimulated by shot noise (a flow of random spikes bombarding neurons). Using an exactly solvable cortical model of neuronal networks on classical random networks, we find critical phenomena accompanying the transitions and their dependence on the shot noise intensity. We show that a pattern of spontaneous neuronal activity near a critical point of a phase transition is a characteristic property that can be used to identify the bifurcation mechanism of the transition. We demonstrate that bursts …
There is growing evidence in favour of the temporal-coding hypothesis that temporal correlation of neuronal discharges may serve to bind distributed neuronal activity into unique representations and, in particular, that $θ$ (3.5-7.5 Hz) and $δ$ ($0.5<$3.5 Hz) oscillations facilitate information coding. The $θ$ and $δ$ rhythms are shown to be involved in various sleep stages, and during anæsthesia, and they undergo changes with the depth of anæsthesia. We introduce a thalamocortical model of interacting neuronal ensembles to describe phase relationships between $θ$ and $δ$ oscillations, especia…
We establish a general linear response relation for spiking neuronal networks, based on chains with unbounded memory. This relation allows us to predict the influence of a weak amplitude time-dependent external stimuli on spatio-temporal spike correlations, from the spontaneous statistics (without stimulus) in a general context where the memory in spike dynamics can extend arbitrarily far in the past. Using this approach, we show how linear response is explicitly related to neuronal dynamics with an example, the gIF model, introduced by M. Rudolph and A. Destexhe. This example illustrates the …
Synchronized bursts (SBs) with complex structures are common in neuronal cultures. Although the origin of SBs is still unclear, they have been studied for their information processing capabilities. Here, we investigate the properties of these SBs in a culture on multi-electrode array system. We find that structures of these SBs are related to the different developmental stages of the cultures. A model based on short term synaptic plasticity, recurrent connections and astrocytic recycling of neurotransmitters has been developed successfully to understand these structures. A phase diagram obtain…
The Kuramoto model (KM) is a theoretical paradigm for investigating the emergence of rhythmic activity in large populations of oscillators. A remarkable example of rhythmogenesis is the feedback loop between excitatory (E) and inhibitory (I) cells in large neuronal networks. Yet, although the EI-feedback mechanism plays a central role in the generation of brain oscillations, it remains unexplored whether the KM has enough biological realism to describe it. Here we derive a two-population KM that fully accounts for the onset of EI-based neuronal rhythms and that, as the original KM, is analytic…
Cortical regions without direct neuronal connections have been observed to exhibit synchronized dynamics. A recent empirical study has further revealed that such regions that share more common neighbors are more likely to behave coherently. To analytically investigate the underlying mechanisms, we consider that a set of n oscillators, which have no direct connections, are linked through m intermediate oscillators (called mediators), forming a complete bipartite network structure. Modeling the oscillators by the Kuramoto-Sakaguchi model, we rigorously prove that mediated remote synchronization,…
Neurons in many brain areas can develop pronounced depolarized state of membrane potential (up state) in addition to the normal hyperpolarized down state near the resting potential. The influence of the up state on signal encoding, however, is not well investigated. Here we construct a one-dimensional bistable neuron model and calculate the linear response to noisy oscillatory inputs analytically. We find that with the appearance of an up state, the transmission function is enhanced by the emergence of a local maximum at some optimal frequency and the phase lag relative to the input signal is …
We study the reliability of large networks of coupled neural oscillators in response to fluctuating stimuli. Reliability means that a stimulus elicits essentially identical responses upon repeated presentations. We view the problem on two scales: neuronal reliability, which concerns the repeatability of spike times of individual neurons embedded within a network, and pooled-response reliability, which addresses the repeatability of the total synaptic output from the network. We find that individual embedded neurons can be reliable or unreliable depending on network conditions, whereas pooled r…
Hysteresis and multistability in network spreading with neuronal activity feedback
Wu Tsai Institute, Yale University（耶鲁大学吴-tsai研究所）；Department of Psychology, Yale University（耶鲁大学心理学系）；Department of Biomedical Engineering, Yale University（耶鲁大学生物医学工程系）
A model of the columnar functional organization of neocortical association areas is studied. The neuronal network is composed of many Hebbian autoassociators, or modules, each of which interacts with a relatively small number of the others. Every module encodes and stores a number of elementary percepts, or features. Memory items, or patterns, are peculiar combinations of features sparsely distributed over the multi-modular network. Any feature stored in any module can be involved in several of the stored patterns; feature-sharing is in fact source of local ambiguities and, consequently, a pot…
Sleep is essential for the maintenance of the brain and the body, yet many features of sleep are poorly understood and mathematical models are an important tool for probing proposed biological mechanisms. The most well-known mathematical model of sleep regulation, the two-process model, models the sleep-wake cycle by two oscillators: a circadian oscillator and a homeostatic oscillator. An alternative, more recent, model considers the mutual inhibition of sleep promoting neurons and the ascending arousal system regulated by homeostatic and circadian processes. Here we show there are fundamental…
The result of computational operations performed at the single cell level are coded into sequences of action potentials (APs). In the cerebral cortex, due to its columnar organization, large number of neurons are involved in any individual processing task. It is therefore important to understand how the properties of coding at the level of neuronal populations are determined by the dynamics of single neuron AP generation. Here we analyze how the AP generating mechanism determines the speed with which an ensemble of neurons can represent transient stochastic input signals. We analyze a generali…
We present a use of modern data-based machine learning approaches to suppress self-sustained collective oscillations typically signaled by ensembles of degenerative neurons in the brain. The proposed hybrid model relies on two major components: an environment of oscillators and a policy-based reinforcement learning block. We report a model-agnostic synchrony control based on proximal policy optimization and two artificial neural networks in an Actor-Critic configuration. A class of physically meaningful reward functions enabling the suppression of collective oscillatory mode is proposed. The s…
We show that for two identical neuronal oscillators with strictly positive phase resetting curve, isochronous synchrony is an unstable attractor and arbitrarily weak noise can destroy entrainment and generate intermittent phase slips. Small inhomogeneity (mismatch in the intrinsic firing rate of the neurons) can stabilize the phase locking and lead to more precise relative spike timing of the two neurons. The results can explain how for a class of neuronal models, including leaky integrate-fire model, inhomogeneity can increase correlation of spike trains when the neurons are synaptically conn…
In this paper, we introduce and systematically study, in terms of phase response curves (PRC), the effect of a dual pulse excitation on the dynamics of an autonomous oscillator. Specifically, we test the deviations from a linear summation of phase advances from two small perturbations. We derive analytically a correction term, which generally appears for oscillators whose intrinsic dimensionality is greater than one. We demonstrate this effect in case of the Stuart-Landau model, and also in various higher dimensional neuronal model. The non-linear correction is found to be proportional to the …
Motivated by recent observations in neuronal systems we investigate all-to-all networks of non-identical oscillators with adaptive coupling. The adaptation models spike-timing-dependent plasticity in which the sum of the weights of all incoming links is conserved. We find multiple phase-locked states that fall into two classes: near-synchronized states and splay states. Among the near-synchronized states are states that oscillate with a frequency that depends only very weakly on the coupling strength and is essentially given by the frequency of one of the oscillators, which is, however, neithe…
We find that sensory noise delivered together with a weak periodic signal not only enhances nonlinear response of neuronal networks, but also improves the synchronization of the response to the signal. We reveal this phenomenon in neuronal networks that are in a dynamical state near a saddle-node bifurcation corresponding to appearance of sustained network oscillations. In this state, even a weak periodic signal can evoke sharp nonlinear oscillations of neuronal activity. These sharp network oscillations have a deterministic form and amplitude determined by nonlinear dynamical equations. The s…
Intracellular access with high spatiotemporal resolution can enhance our understanding of how neurons or cardiomyocytes regulate and orchestrate network activity, and how this activity can be affected with pharmacology or other interventional modalities. Nanoscale devices often employ electroporation to transiently permeate the cell membrane and record intracellular potentials, which tend to decrease rapidly to extracellular potential amplitudes with time. Here, we report innovative scalable, vertical, ultra-sharp nanowire arrays that are individually addressable to enable long-term, native re…
Synaptic delays modulate population phase and amplitude responses in oscillatory excitatory-inhibitory networks
Institute for Advanced Studies in Basic Sciences (IASBS)（基础科学高等研究院）；Institute for Research in Fundamental Sciences (IPM)（基本科学研究院）；Pasargad Institute for Advanced Innovative Solutions (PIAIS)（帕萨尔加德先进创新解决方案研究所）；The Zapata-Briceño Institute of Neuroscience（萨帕塔-布里斯尼奥神经科学研究所）
We consider a simple Markovian class of the stochastic Wilson-Cowan type models of neuronal network dynamics, which incorporates stochastic delay caused by the existence of a refractory period of neurons. From the point of view of the dynamics of the individual elements, we are dealing with a network of non-Markovian stochastic two-state oscillators with memory which are coupled globally in a mean-field fashion. This interrelation of a higher-dimensional Markovian and lower-dimensional non-Markovian dynamics is discussed in its relevance to the general problem of the network dynamics of comple…
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…
Information transmission is a key element for information processing in the brain. A number of mechanisms have been proposed for transferring volleys of spikes between layers of a feedforward neural circuit. Many of these mechanisms use synchronous activity to provide windows in time when spikes may be transferred more easily from layer to layer. Recently, we have demonstrated that a pulse-gating mechanism can transfer graded information between layers in a feedforward neuronal network. Our transfer mechanism resulted in a time-translationally invariant firing rate and synaptic current wavefor…
Persistent activity is postulated to drive neural network plasticity and learning. To investigate its underlying cellular mechanisms, we developed a biophysically tractable model that explains the emergence, sustenance, and eventual termination of short-term persistent activity. Using the model, we reproduced the features of reverberating activity that were observed in small (50-100 cells) networks of cultured hippocampal neurons, such as the appearance of polysynaptic current clusters, the typical inter-cluster intervals, the typical duration of reverberation, and the response to changes in e…
Activity in coupled systems is often oscillatory, for example, the firing pattern of neuronal populations. Whereas these oscillations have been studied predominantly in local circuits, here we show how the topology of large-scale networks, leading to large feedback loops, influences oscillations in the resting state. We find that the hierarchical modular organization of neuronal networks supports distinct spectral patterns of neural rhythms similar to those observed experimentally in different species such as rat and human. For individual neurons, multiple peak frequencies with non-integer rat…
We have identified the occurrence of chimera states for various coupling schemes in networks of two-dimensional and three-dimensional Hindmarsh-Rose oscillators, which represent realistic models of neuronal ensembles. This result, together with recent studies on multiple chimera states in nonlocally coupled FitzHugh-Nagumo oscillators, provide strong evidence that the phenomenon of chimeras may indeed be relevant in neuroscience applications. Moreover, our work verifies the existence of chimera states in coupled bistable elements, whereas to date chimeras were known to arise in models possessi…
