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 and avalanches are precursors of a first-order phase transition, paroxysmal-like spikes of activity precede a second-order phase transition caused by a saddle-node bifurcation, while irregular spindle oscillations represent spontaneous activity near a second-order phase transition caused by a supercritical Hopf bifurcation. Our most interesting result is the observation of the paroxysmal-like spikes. We show that a paroxysmal-like spike is a single nonlinear event that appears instantly from a low background activity with a rapid onset, reaches a large amplitude, and ends up with an abrupt return to lower activity. These spikes are similar to single paroxysmal spikes and sharp waves observed in EEG measurements. Our analysis shows that above the saddle-node bifurcation, sustained network oscillations appear with a large amplitude but a small frequency in contrast to network oscillations near the Hopf bifurcation that have a small amplitude but a large frequency. We discuss an amazing similarity between excitability of the cortical model stimulated by shot noise and excitability of the Morris-Lecar neuron stimulated by an applied current.
Noise is an inherent part of neuronal dynamics, and thus of the brain. It can be observed in neuronal activity at different spatiotemporal scales, including in neuronal membrane potentials, local field potentials, electroencephalography, and magnetoencephalography. A central research topic in contemporary neuroscience is to elucidate the functional role of noise in neuronal information processing. Experimental studies have shown that a suitable level of noise may enhance the detection of weak neuronal signals by means of stochastic resonance. In response, theoretical research, based on the the…
Noise appears in the brain due to various sources, such as ionic channel fluctuations and synaptic events. They affect the activities of the brain and influence neuron action potentials. Stochastic differential equations have been used to model firing patterns of neurons subject to noise. In this work, we consider perturbing noise in the adaptive exponential integrate-and-fire (AEIF) neuron. The AEIF is a two-dimensional model that describes different neuronal firing patterns by varying its parameters. Noise is added in the equation related to the membrane potential. We show that a noise curre…
Using an exactly solvable cortical model of a neuronal network, we show that, by increasing the intensity of shot noise (flow of random spikes bombarding neurons), the network undergoes first- and second-order non-equilibrium phase transitions. We study the nature of the transitions, bursts and avalanches of neuronal activity. Saddle-node and supercritical Hopf bifurcations are the mechanisms of emergence of sustained network oscillations. We show that the network stimulated by shot noise behaves similar to the Morris-Lecar model of a biological neuron stimulated by an applied current.
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…
Integrated Information Theory (IIT) is an audacious attempt to pin down the abstract, phenomenological experiences of consciousness into a rigorous, mathematical framework. We show that IIT's stance in regards to neuronal noise is inconsistent with experimental data demonstrating that neuronal noise in the brain plays a critical role in learning, visual recognition, and even categorical representation. IIT predicts that entropy due to noise will reduce the information integration of a physical system, which is inconsistent with experimental data demonstrating that decision-related noise is a n…
Spontaneous formation of clusters of synchronized spiking in a structureless ensemble of equal stochastically perturbed excitable neurons with delayed coupling is demonstrated for the first time. The effect is a consequence of a subtle interplay between interaction delays, noise and the excitable character of a single neuron. Dependence of the cluster properties on the time-lag, noise intensity and the synaptic strength is investigated.
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 …
First spike latency following stimulus onset is of significant physiological relevance. Neurons transmit information about their inputs by transforming them into spike trains, and the timing of these spike trains is in turn crucial for effectively encoding that information. Random processes and uncertainty that underly neuronal dynamics have been shown to prolong the time towards the first response in a phenomenon dubbed noise-delayed decay. Here we study whether Hodgkin-Huxley neurons with a tunable intensity of intrinsic noise might have shorter response times to external stimuli just above …
Avalanches with power-law distributed size parameters have been observed in neuronal networks. This observation might be a manifestation of the self-organized criticality (SOC). Yet, the physiological mechanicsm of this behavior is currently unknown. Describing synaptic noise as transmission failures mainly originating from the probabilistic nature of neurotransmitter release, this study investigates the potential of this noise as a mechanism for driving the functional architecture of the neuronal networks towards SOC. To this end, a simple finite state neuron model, with activity dependent an…
We study the role of scale-free structure and noise in collective dynamics of neuronal networks. For this purpose, we simulate and study analytically a cortical circuit model with stochastic neurons. We compare collective neuronal activity of networks with different topologies: classical random graphs and scale-free networks. We show that, in scale-free networks with divergent second moment of degree distribution, an influence of noise on neuronal activity is strongly enhanced in comparison with networks with a finite second moment. A very small noise level can stimulate spontaneous activity o…
Harmonic fractal transformation for modeling complex neuronal effects: from bursting and noise shaping to waveform sensitivity and noise-induced subthreshold spiking
Neurons in the central nervous system are affected by complex and noisy signals due to fluctuations in their cellular environment and in the inputs they receive from many other cells 1,2. Such noise usually increases the probability that a neuron will send out a signal to its target cells 2-5. In stochastic resonance, which occurs in many physical and biological systems, an optimal response is found at a particular noise amplitude 6-9. We have found that in a classical neuronal model the opposite can occur - that noise can subdue or turn off repetitive neuronal activity in both single cells an…
Biological systems are notoriously noisy. Noise, therefore, also plays an important role in many models of neural impulse generation. Noise is not only introduced for more realistic simulations but also to account for cooperative effects between noisy and nonlinear dynamics. Often, this is achieved by a simple noise term in the membrane equation (current noise). However, there are ongoing discussions whether such current noise is justified or whether rather conductance noise should be introduced because it is closer to the natural origin of noise. Therefore, we have compared the effects of cur…
Neuronal responses to sensory stimuli or neuronal responses related to behaviour are often extracted by averaging neuronal activity over large number of experimental trials. Such trial-averaging is carried out to reduce noise and to reduce the influence of other signals unrelated to the corresponding stimulus or behaviour. However, if the recorded neuronal responses are jittered in time with respect to the corresponding stimulus or behaviour, averaging over trials may distort the estimation of the underlying neuronal response. Here, we present an algorithm, named dTAV algorithm, for realigning…
Recent experiments have shown that the spontaneous activity of young dissociated neuronal cultures can be described as a process of highly inhomogeneous nucleation and front propagation due to the localization of noise activity, i.e., noise focusing. However, the basic understanding of the mechanisms of noise build-up leading to the nucleation remain an open fundamental problem. Here we present a minimal dynamical model called stochastic quorum percolation that can account for the observed phenomena, while providing a robust theoretical framework. The model reproduces the first and second orde…
We consider signal transaction in a simple neuronal model featuring intrinsic noise. The presence of noise limits the precision of neural responses and impacts the quality of neural signal transduction. We assess the signal transduction quality in relation to the level of noise, and show it to be maximized by a non-zero level of noise, analogous to the stochastic resonance effect. The quality enhancement occurs for a finite range of stimuli to a single neuron; we show how to construct networks of neurons that extend the range. The range increases more rapidly with network size when we make use…
Recurrent networks of dynamic elements frequently exhibit emergent collective oscillations, which can display substantial regularity even when the individual elements are considerably noisy. How noise-induced dynamics at the local level coexists with regular oscillations at the global level is still unclear. Here we show that a combination of stochastic recurrence-based initiation with deterministic refractoriness in an excitable network can reconcile these two features, leading to maximum collective coherence for an intermediate noise level. We report this behavior in the slow oscillation reg…
We study the characteristics of weak signal detection by a recurrent neuronal network with plastic synaptic coupling. It is shown that in the presence of an asynchronous component in synaptic transmission, the network acquires selectivity with respect to the frequency of weak periodic stimuli. For non-periodic frequency-modulated stimuli, the response is quantified by the mutual information between input (signal) and output (network's activity), and is optimized by synaptic depression. Introducing correlations in signal structure resulted in the decrease of input-output mutual information. Our…
We demonstrate that human electrophysiological recordings of the local field potential (LFP) from intracranial electrodes, acquired from a variety of cerebral regions, show a ubiquitous $1/f^2$ scaling within the power spectrum. We develop a quantitative model that treats the generation of these fields in an analogous way to that of electronic shot noise, and use this model to specifically address the cause of this $1/f^2$ Brownian noise. The model gives way to two analytically tractable solutions, both displaying Brownian noise: 1) uncorrelated cells that display sharp initial activity, whose…
Filtered shot noise processes have proven to be very effective in modelling the evolution of systems exposed to stochastic shot noise sources, and have been applied to a wide variety of fields ranging from electronics through biology. In particular, they can model the membrane potential Vm of neurons driven by stochastic input, where these filtered processes are able to capture the non-stationary characteristics of Vm fluctuations in response to pre-synaptic input with variable rate. In this paper, we apply the general framework of Poisson Point Processes transformations to analyse these syste…
In this paper, we systematically investigate both the synfire propagation and firing rate propagation in feedforward neuronal network coupled in an all-to-all fashion. In contrast to most earlier work, where only reliable synaptic connections are considered, we mainly examine the effects of unreliable synapses on both types of neural activity propagation in this work. We first study networks composed of purely excitatory neurons. Our results show that both the successful transmission probability and excitatory synaptic strength largely influence the propagation of these two types of neural act…
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 …
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…
The stochastic opening and closing of voltage-gated ion channels produces noise in neurons. The effect of this noise on the neuronal performance has been modelled using either approximate or Langevin model, based on stochastic differential equations or an exact model, based on a Markov process model of channel gating. Yet whether the Langevin model accurately reproduces the channel noise produced by the Markov model remains unclear. Here we present a comparison between Langevin and Markov models of channel noise in neurons using single compartment Hodgkin-Huxley models containing either $Na^{+…
A new class of energy-efficient digital microprocessor is being developed which is susceptible to thermal noise and consequently operates in probabilistic rather than conventional deterministic mode. Hybrid computing systems which combine probabilistic and deterministic processors can provide robust and efficient tools for computational problems that hitherto would be intractable by conventional deterministic algorithm. These developments suggest a revised perspective on the consequences of ion-channel noise in slender axons, often regarded as a hindrance to neuronal computations. It is propos…
In a generic neuron model, we present the linear response theory for the firing rate in response to both time dependent input currents and noise amplitudes. In both cases the signal transmission is strongly attenuated for frequencies above the stationary firing rate. For high frequencies both the mean input and the noise transmission function decay as ω^{-2}, independent of model details. Our results indicate that previously suggested mechanisms for near instantaneous transmission of information are not consistent with the spike generation mechanism of real neurons.
An appealing new principle for neural population codes is that correlations among neurons organize neural activity patterns into a discrete set of clusters, which can each be viewed as a noise-robust population "codeword". Previous studies assumed that these codewords corresponded geometrically with local peaks in the probability landscape of neural population responses. Here, we analyze multiple datasets of the responses of ~150 retinal ganglion cells and show that local probability peaks are absent under broad, non-repeated stimulus ensembles, which are characteristic of natural behavior. Ho…
Many systems in Nature exhibit avalanche dynamics with scale-free features. A general scaling theory has been proposed for critical avalanche profiles in crackling noise, predicting the collapse onto a universal avalanche shape, as well as the scaling behaviour of the activity power spectrum as Brown noise. Recently, much attention has been given to the profile of neuronal avalanches, measured in neuronal systems in vitro and in vivo. Although a universal profile was evidenced, confirming the validity of the general scaling theory, the parallel study of the power spectrum scaling under the sam…
The threshold voltage for action potential generation is a key regulator of neuronal signal transduction, yet the mechanism of its dynamic variation is still not well described. In this paper, we propose that threshold phenomena can be classified as parameter thresholds and state thresholds. Voltage thresholds which belong to the state threshold are determined by the `general separatrix' in state space. We demonstrate that the separatrix generally exists in the state space of neuron models. The general form of separatrix was assumed as the function of both states and stimuli and the previously…
The output signal is examined for the Jacobi neuronal model which is characterized by input-dependent multiplicative noise. The dependence of the noise on the rate of inhibition turns out to be of primary importance to observe maxima both in the output firing rate and in the diffusion coefficient of the spike count and, simultaneously, a minimum in the coefficient of variation (Fano factor). Moreover, we observe that an increment of the rate of inhibition can increase the degree of coherence computed from the power spectrum. This means that inhibition can enhance the coherence and thus the inf…
We consider an excitatory population of subthreshold Izhikevich neurons which cannot fire spontaneously without noise. As the coupling strength passes a threshold, individual neurons exhibit noise-induced burstings. This neuronal population has adaptive dynamic synaptic strengths governed by the spike-timing-dependent plasticity (STDP). In the absence of STDP, stochastic burst synchronization (SBS) between noise-induced burstings of subthreshold neurons was previously found to occur over a large range of intermediate noise intensities. Here, we study the effect of additive STDP on the SBS by v…
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…
Stochastic dynamics and synchronization in motif-based neuronal networks
Toronto Metropolitan University（多伦多都会大学）；University of Toronto（多伦多大学）；Florida State University（佛罗里达州立大学）；Institute of Molecular Biophysics, Florida State University（佛罗里达州立大学分子生物物理研究所）；St. Michael’s Hospital, Unity Health Toronto（圣迈克尔医院（联合健康多伦多））
We investigate front propagation and synchronization transitions in dependence on the information transmission delay and coupling strength over scale-free neuronal networks with different average degrees and scaling exponents. As the underlying model of neuronal dynamics, we use the efficient Rulkov map with additive noise. We show that increasing the coupling strength enhances synchronization monotonously, whereas delay plays a more subtle role. In particular, we found that depending on the inherent oscillation frequency of individual neurons, regions of irregular and regular propagating exci…
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 two recent papers, Rudolph and Destexhe (Neural Comp. {\bf 15}, 2577-2618, 2003; Neural Comp. in press, 2005) studied a leaky integrator model (i.e. an RC-circuit) driven by correlated (``colored'') Gaussian conductance noise and Gaussian current noise. In the first paper they derived an expression for the stationary probability density of the membrane voltage; in the second paper this expression was modified to cover a larger parameter regime. Here we show by standard analysis of solvable limit cases (white-noise limit of additive and multiplicative noise sources; only slow multiplicative …
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…
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…
We investigate the synchronization properties between two excitatory coupled neurons in the presence of an inhibitory loop mediated by an interneuron. Dynamical inhibition together with noise independently applied to each neuron provide phase diversity in the dynamics of the neuronal motif. We show that the interplay between the coupling strengths and external noise controls the phase relations between the neurons in a counter-intuitive way. For a master-slave configuration (unidirectional coupling) we find that the slave can anticipate the master, on average, if the slave is subject to the in…
Subjective tinnitus (ST) is generally assumed to be a consequence of hearing loss (HL). In animal studies acoustic trauma can lead to behavioral signs of ST, in human studies ST patients without increased hearing thresholds were found to suffer from so called hidden HL. Additionally, ST is correlated with pathologically increased spontaneous firing rates and neuronal hyperactivity (NH) along the auditory pathway. Homeostatic plasticity (HP) has been proposed as a compensation mechanism leading to the development of NH, arguing that after HL initially decreased mean firing rates of neurons are …
Identifying the right tools to express the stochastic aspects of neural activity has proven to be one of the biggest challenges in computational neuroscience. Even if there is no definitive answer to this issue, the most common procedure to express this randomness is the use of stochastic models. In accordance with the origin of variability, the sources of randomness are classified as intrinsic or extrinsic and give rise to distinct mathematical frameworks to track down the dynamics of the cell. While the external variability is generally treated by the use of a Wiener process in models such a…
Neuronal systems have been modeled by complex networks in different description levels. Recently, it has been verified that networks can simultaneously exhibit one coherent and other incoherent domain, known as chimera states. In this work, we study the existence of chimera states in a network considering the connectivity matrix based on the cat cerebral cortex. The cerebral cortex of the cat can be separated in 65 cortical areas organised into the four cognitive regions: visual, auditory, somatosensory-motor and frontolimbic. We consider a network where the local dynamics is given by the Hind…
We numerically investigate the influence of intrinsic channel noise on the dynamical response of delay-coupling in neuronal systems. The stochastic dynamics of the spiking is modeled within a stochastic modification of the standard Hodgkin-Huxley model wherein the delay-coupling accounts for the finite propagation time of an action potential along the neuronal axon. We quantify this delay-coupling of the Pyragas-type in terms of the difference between corresponding presynaptic and postsynaptic membrane potentials. For an elementary neuronal network consisting of two coupled neurons we detect c…
Here, we consider the open issue of how the energy efficiency of neural information transmission process in a general neuronal array constrains the network size, and how well this network size ensures the neural information being transmitted reliably in a noisy environment. By direct mathematical analysis, we have obtained general solutions proving that there exists an optimal neuronal number in the network with which the average coding energy cost (defined as energy consumption divided by mutual information) per neuron passes through a global minimum for both subthreshold and superthreshold s…
Two dynamical systems unidirectionally coupled in a sender-receiver configuration can synchronize with a nonzero phase-lag. In particular, the system can exhibit anticipated synchronization (AS), which is characterized by a negative phase-lag, if the receiver (R) also receives a delayed negative self-feedback. Recently, AS was shown to occur between cortical-like neuronal populations in which the self-feedback is mediated by inhibitory synapses. In this biologically plausible scenario, a transition from the usual delayed synchronization (DS, with positive phase-lag) to AS can be mediated by th…
Biocomputing technologies exploit biological communication mechanisms involving cell-cell signal propagation to perform computations. Researchers recently worked toward realising logic gates made by neurons to develop novel devices such as organic neuroprostheses or brain implants made by cells, herein termed living implants. Several challenges arise from this approach, mainly associated with the stochastic nature and noise of neuronal communication. Since astrocytes play a crucial role in the regulation of neurons activity, there is a possibility whereby astrocytes can be engineered to contro…
Brain plasticity, also known as neuroplasticity, is a fundamental mechanism of neuronal adaptation in response to changes in the environment or due to brain injury. In this review, we show our results about the effects of synaptic plasticity on neuronal networks composed by Hodgkin-Huxley neurons. We show that the final topology of the evolved network depends crucially on the ratio between the strengths of the inhibitory and excitatory synapses. Excitation of the same order of inhibition revels an evolved network that presents the rich-club phenomenon, well known to exist in the brain. For ini…
Spontaneous neuronal activity is a ubiquitous feature of cortex. Its spatiotemporal organization reflects past input and modulates future network output. Here we study whether a particular type of spontaneous activity is generated by a network that is optimized for input processing. Neuronal avalanches are a type of spontaneous activity observed in superficial cortical layers in vitro and in vivo with statistical properties expected from a network in a 'critical state'. Theory predicts that the critical state and, therefore, neuronal avalanches are optimal for input processing, but until now, …
