arXiv subjects

Alessandro Torcini

Publications and source records attributed to Alessandro Torcini.

At least 19 recordsLinked to original sources

arXiv2025

A theory for self-sustained balanced states in absence of strong external currents

Recurrent neural networks with balanced excitation and inhibition exhibit irregular asynchronous dynamics, which is fundamental for cortical computations. Classical balance mechanisms require strong external inputs to sustain finite firing rates, raising concerns about their biological plausibility. Here, we investigate an alternative mechanism based on short-term synaptic depression (STD) acting on excitatory-excitatory synapses, which dynamically balances the network activity without the need of external inputs. By employing numerical simulations and theoretical investigations we characterize the dynamics of a massively coupled network made up of $N$ rate-neuron models. Depending on the synaptic strength $J_0$, the network exhibits two distinct regimes: at sufficiently small $J_0$, it converges to a homogeneous fixed point, while for sufficiently large $J_0$, it exhibits Rate Chaos. For finite networks, we observe several different routes to chaos depending on the network realization. The width of the transition region separating the homogeneous stable solution from Rate Chaos appears to shrink for increasing $N$ and eventually to vanish in the thermodynamic limit. The characterization of the Rate Chaos regime performed by employing Dynamical Mean Field approaches allow us on one side to confirm that this novel balancing mechanism is able to sustain finite irregular activity even in the thermodynamic limit, and on the other side to reveal that the balancing occurs via dynamic cancellation of the input correlations generated by the massive coupling. Our findings show that STD provides an intrinsic self-regulating mechanism for balanced networks, sustaining irregular yet stable activity without the need of biologically unrealistic inputs. This work extends the balanced network paradigm, offering insights into how cortical circuits could maintain robust dynamics via synaptic adaptation.

q-bio.NC↗

arXiv2025

Is the cortical dynamics ergodic? A numerical study in partially symmetric networks of spiking neurons

Cortical activity in-vivo displays relaxational time scales much longer than the membrane time constant of the neurons or the deactivation time of ionotropic synaptic conductances. The mechanisms responsible for such slow dynamics are not understood. Here, we show that slow dynamics naturally and robustly emerges in dynamically-balanced networks of spiking neurons. This requires only partial symmetry in the synaptic connectivity, a feature of local cortical networks observed in experiments. The symmetry generates an effective, excitatory self-coupling of the neurons that leads to long-lived fluctuations in the network activity, without destroying the dynamical balance. When the excitatory self-coupling is suitably strong, the same mechanism leads to multiple equilibrium states of the network dynamics. Our results reveal a novel dynamical regime of the collective activity in spiking networks, where the memory of the initial state persists for very long times and ergodicity is broken.

cond-mat.dis-nn↗

arXiv2025

Synaptic shot-noise triggers fast and slow global oscillations in balanced neural networks

Neural dynamics is determined by the transmission of discrete synaptic pulses (synaptic shot-noise) among neurons. However, the neural responses are usually obtained within the diffusion approximation modeling synaptic inputs as continuous Gaussian noise. Here, we present a rigorous mean-field theory that encompasses synaptic shot-noise for sparse balanced inhibitory neural networks driven by an excitatory drive. Our theory predicts new dynamical regimes, in agreement with numerical simulations, which are not captured by the classical diffusion approximation. Notably, these regimes feature self-sustained global oscillations emerging at low connectivity (in-degree) via either continuous or hysteretic transitions and characterized by irregular neural activity, as expected for balanced dynamics. For sufficiently weak (strong) excitatory drive (inhibitory feedback) the transition line displays a peculiar re-entrant shape revealing the existence of global oscillations at low and high in-degrees, separated by an asynchronous regime at intermediate levels of connectivity. The mechanisms leading to the emergence of these global oscillations are distinct: drift-driven at high connectivity and cluster activation at low connectivity. The frequency of these two kinds of global oscillations can be varied from slow (around 1 Hz) to fast (around 100 Hz), without altering their microscopic and macroscopic features, by adjusting the excitatory drive and the synaptic inhibition strength in a prescribed way. Furthermore, the cluster-activated oscillations at low in-degrees could correspond to the gamma rhythms reported in mammalian cortex and hippocampus and attributed to ensembles of inhibitory neurons sharing few synaptic connections [G. Buzsaki and X.-J. Wang, Annual Review of Neuroscience 35, 203 (2012)].

cond-mat.dis-nn↗

arXiv2024

Chaotic synchronization in adaptive networks of pulse-coupled oscillators

Ensembles of phase-oscillators are known to exhibit a variety of collective regimes. Here, we show that a simple mean-field model involving two heterogenous populations of pulse-coupled oscillators, exhibits, in the strong-coupling limit, a robust irregular macroscopic dynamics. The resulting, strongly synchronized, regime is sustained by a homeostatic mechanism induced by the shape of the phase-response curve combined with adaptive coupling strength, included to account for energy dissipated by the pulse emission. The proposed setup mimicks a neural network composed of excitatory and inhibitory neurons.

cond-mat.dis-nn↗

arXiv2024

Emergence and maintenance of modularity in neural networks with Hebbian and anti-Hebbian inhibitory STDP

The modular and hierarchical organization of the brain is believed to support the coexistence of segregated (specialization) and integrated (binding) information processes. A relevant question is yet to understand how such architecture naturally emerges and is sustained over time, given the plastic nature of the brain's wiring. Following evidences that the sensory cortices organize into assemblies under selective stimuli, it has been shown that stable neuronal assemblies can emerge due to targeted stimulation, embedding various forms of synaptic plasticity in presence of homeostatic and/or control mechanisms. Here, we show that simple spike-timing-dependent plasticity (STDP) rules, based only on pre- and post-synaptic spike times, can also lead to the stable encoding of memories in the absence of any control mechanism. We develop a model of spiking neurons, trained by stimuli targeting different subpopulations. The model satisfies some biologically plausible features: (i) it contains excitatory and inhibitory neurons with Hebbian and anti-Hebbian STDP; (ii) neither the neuronal activity nor the synaptic weights are frozen after the learning phase. Instead, the neurons are allowed to fire spontaneously while synaptic plasticity remains active. We find that only the combination of two inhibitory STDP subpopulations allows for the formation of stable modules in the network, with each subpopulation playing a distinctive role. The Hebbian subpopulation controls for the firing activity, while the anti-Hebbian neurons promote pattern selectivity. After the learning phase, the network settles into an asynchronous irregular resting-state. This post-learning activity is associated with spontaneous memory recalls which turn out to be fundamental for the long-term consolidation of the learned memories.

q-bio.NC↗

arXiv2024

Coexistence of asynchronous and clustered dynamics in noisy inhibitory neural networks

A regime of coexistence of asynchronous and clustered dynamics is analyzed for globally coupled homogeneous and heterogeneous inhibitory networks of quadratic integrate-and-fire (QIF) neurons subject to Gaussian noise. The analysis is based on accurate extensive simulations and complemented by a mean-field description in terms of low-dimensional next generation neural mass models for heterogeneously distributed synaptic couplings. The asynchronous regime is observable at low noise and becomes unstable via a sub-critical Hopf bifurcation at sufficiently large noise. This gives rise to a coexistence region between the asynchronous and the clustered regime. The clustered phase is characterized by population bursts in the {\gamma}-range (30-120 Hz), where neurons are split in two equally populated clusters firing in alternation. This clustering behaviour is quite peculiar: despite the global activity being essentially periodic, single neurons display switching between the two clusters due to heterogeneity and/or noise.

cond-mat.dis-nn↗

arXiv2024

A robust balancing mechanism for spiking neural networks

Dynamical balance of excitation and inhibition is usually invoked to explain the irregular low firing activity observed in the cortex. We propose a robust nonlinear balancing mechanism for a random network of spiking neurons, which works also in absence of strong external currents. Biologically, the mechanism exploits the plasticity of excitatory-excitatory synapses induced by short-term depression. Mathematically, the nonlinear response of the synaptic activity is the key ingredient responsible for the emergence of a stable balanced regime. Our claim is supported by a simple self-consistent analysis accompanied by extensive simulations performed for increasing network sizes. The observed regime is essentially fluctuation driven and characterized by highly irregular spiking dynamics of all neurons.

cond-mat.dis-nn↗

arXiv2023

Discrete synaptic events induce global oscillations in balanced neural networks

Neural dynamics is triggered by discrete synaptic inputs of finite amplitude. However, the neural response is usually obtained within the diffusion approximation (DA) representing the synaptic inputs as Gaussian noise. We derive a mean-field formalism encompassing synaptic shot-noise for sparse balanced networks of spiking neurons. For low (high) external drives (synaptic strengths) irregular global oscillations emerge via continuous and hysteretic transitions, correctly predicted by our approach, but not from the DA. These oscillations display frequencies in biologically relevant bands.

q-bio.NC↗

arXiv2023

Cluster States and $\pi$-Transition in the Kuramoto Model with Higher Order Interactions

We have examined the synchronization and de-synchronization transitions observable in the Kuramoto model with a standard pair-wise first harmonic interaction plus a higher order (triadic) symmetric interaction for unimodal and bimodal Gaussian distributions of the natural frequencies $\{ \omega_i \}$. These transitions have been accurately characterized thanks to a self-consistent mean-field approach joined with extensive numerical simulations. The higher-order interactions favour the formation of two cluster states, which emerge from the incoherent regime via continuous (discontinouos) transitions for unimodal (bimodal) distributions. Fully synchronized initial states give rise to two symmetric equally populated bimodal clusters, each characterized by either positive or negative natural frequencies. These bimodal clusters are formed at an angular distance $\gamma$, which increases for decreasing pair-wise couplings until it reaches $\gamma=\pi$ (corresponding to an anti-phase configuration), where the cluster state destabilizes via an abrupt transition: the $\pi$-transition. The uniform clusters that reform immediately after (with a smaller angle $\gamma$) are composed by oscillators with positive and negative $\{ \omega_i \}$. For bimodal distributions we have obtained detailed phase diagrams involving all the possible dynamical states in terms of standard and novel order parameters. In particular, the clustering order parameter, here introduced, appears quite suitable to characterize the two cluster regime. As a general aspect, hysteretic (non hysteretic) synchronization transitions, mostly mediated by the emergence of standing waves, are observable for attractive (repulsive) higher-order interactions.

nlin.AO↗

arXiv2022

Population spiking and bursting in next generation neural masses with spike-frequency adaptation

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 for the derivation of bifurcation diagrams and the identification of the possible macroscopic regimes emerging both in a single and in two identically coupled neural masses. In single populations SFA favours the emergence of population bursts in excitatory networks, while it hinders tonic population spiking for inhibitory ones. The symmetric coupling of two neural masses, in absence of adaptation, leads to the emergence of macroscopic solutions with broken symmetry: namely, chimera-like solutions in the inhibitory case and anti-phase population spikes in the excitatory one. The addition of SFA leads to new collective dynamical regimes exhibiting cross-frequency coupling (CFC) among the fast synaptic time scale and the slow adaptation one, ranging from anti-phase slow-fast nested oscillations to symmetric and asymmetric bursting phenomena. The analysis of these CFC rhythms in the $\theta$-$\gamma$ range has revealed that a reduction of SFA leads to an increase of the $\theta$ frequency joined to a decrease of the $\gamma$ one. This is analogous to what reported experimentally for the hippocampus and the olfactory cortex of rodents under cholinergic modulation, that is known to reduce SFA.

cond-mat.dis-nn↗

arXiv2022

Collective excitability in highly diluted networks of rotators

We report on collective excitable events in a highly-diluted random network of non-excitable nodes. Excitability arises thanks to a self-sustained local adaptation mechanism that drives the system on a slow time-scale across a hysteretic phase transition involving states with different degrees of synchronization. These phenomena have been investigated for the Kuramoto model with bimodal distribution of the natural frequencies and for the Kuramoto model with inertia and an unimodal frequency distribution. We consider global and local stimulation protocols and characterize the system response for different level of dilution. We compare the results with those obtained in the fully-coupled case showing that such collective phenomena are remarkably robust against network diluteness.

cond-mat.dis-nn↗

arXiv2021

Collective canard explosions of globally-coupled rotators with adaptive coupling

Canards, special trajectories that follow invariant repelling slow manifolds for long time intervals, have been frequently observed in slow-fast systems of either biological, chemical and physical nature. Here, collective canard explosions are demonstrated in a population of globally-coupled phase-rotators subject to adaptive coupling. In particular, we consider a bimodal Kuramoto model displaying coexistence of asynchronous and partially synchronized dynamics subject to a linear global feedback. A detailed geometric singular perturbation analysis of the associated mean-field model allows us to explain the emergence of collective canards in terms of the stability properties of the one-dimensional critical manifold, near which the slow macroscopic dynamics takes place. We finally show how collective canards and related manifolds gradually emerge in the globally-coupled system for increasing system sizes, in spite of the trivial dynamics of the uncoupled rotators.

nlin.AO↗

arXiv2021

Coherent oscillations in balanced neural networks driven by endogenous fluctuations

We present a detailed analysis of the dynamical regimes observed in a balanced network of identical Quadratic Integrate-and-Fire (QIF) neurons with a sparse connectivity for homogeneous and heterogeneous in-degree distribution. Depending on the parameter values, either an asynchronous regime or periodic oscillations spontaneously emerge. Numerical simulations are compared with a mean field model based on a self-consistent Fokker-Planck equation (FPE). The FPE reproduces quite well the asynchronous dynamics in the homogeneous case by either assuming a Poissonian or renewal distribution for the incoming spike trains. An exact self consistent solution for the mean firing rate obtained in the limit of infinite in-degree allows identifying balanced regimes that can be either mean- or fluctuation-driven. A low-dimensional reduction of the FPE in terms of circular cumulants is also considered. Two cumulants suffice to reproduce the transition scenario observed in the network. The emergence of periodic collective oscillations is well captured both in the homogeneous and heterogeneous setups by the mean field models upon tuning either the connectivity, or the input DC current. In the heterogeneous situation we analyze also the role of structural heterogeneity.

q-bio.NC↗

arXiv2021

Asynchronous and coherent dynamics in balanced excitatory-inhibitory spiking networks

Dynamic excitatory-inhibitory (E-I) balance is a paradigmatic mechanism invoked to explain the irregular low firing activity observed in the cortex. However, we will show that the E-I balance can be at the origin of other regimes observable in the brain. The analysis is performed by combining simulations of sparse E-I networks composed of N spiking neurons with analytical investigations of low dimensional neural mass models. The bifurcation diagrams, derived for the neural mass model, allow to classify the asynchronous and coherent behaviours emerging any finite in-degree K. In the limit N >> K >> 1 both supra and sub-threshold balanced asynchronous regimes can be observed. Due to structural heterogeneity the asynchronous states are characterized by the splitting of the neurons in three groups: silent, fluctuation and mean driven. The coherent rhythms are characterized by regular or irregular temporal fluctuations joined to spatial coherence similar to coherent fluctuations observed in the cortex over multiple spatial scales. Collective Oscillations (COs) can emerge due to two different mechanisms. A first mechanism similar to the pyramidal-interneuron gamma (PING) one. The second mechanism is intimately related to the presence of current fluctuations, which sustain COs characterized by an essentially simultaneous bursting of the two populations. We observe period-doubling cascades involving the PING-like COs finally leading to the appearance of coherent chaos. For sufficiently strong current fluctuations we report a novel mechanism of frequency locking among collective rhythms promoted by these intrinsic fluctuations. Our analysis suggest that despite PING-like or fluctuation driven COS are observable for any finite in-degree K, in the limit N >> K >> 1 these solutions result in two coexisting balanced regimes: an asynchronous and a fully synchronized one.

q-bio.NC↗

arXiv2021

A reduction methodology for fluctuation driven population dynamics

Lorentzian distributions have been largely employed in statistical mechanics to obtain exact results for heterogeneous systems. Analytic continuation of these results is impossible even for slightly deformed Lorentzian distributions, due to the divergence of all the moments (cumulants). We have solved this problem by introducing a `pseudo-cumulants' expansion. This allows us to develop a reduction methodology for heterogeneous spiking neural networks subject to extrinsinc and endogenous noise sources, thus generalizing the mean-field formulation introduced in [E. Montbri\'o et al., Phys. Rev. X 5, 021028 (2015)].

cond-mat.stat-mech↗

arXiv2020

Exact neural mass model for synaptic-based working memory

A synaptic theory of Working Memory (WM) has been developed in the last decade as a possible alternative to the persistent spiking paradigm. In this context, we have developed a neural mass model able to reproduce exactly the dynamics of heterogeneous spiking neural networks encompassing realistic cellular mechanisms for short-term synaptic plasticity. This population model reproduces the macroscopic dynamics of the network in terms of the firing rate and the mean membrane potential. The latter quantity allows us to get insight on Local Field Potential and electroencephalographic signals measured during WM tasks to characterize the brain activity. More specifically synaptic facilitation and depression integrate each other to efficiently mimic WM operations via either synaptic reactivation or persistent activity. Memory access and loading are associated to stimulus-locked transient oscillations followed by a steady-state activity in the $\beta-\gamma$ band, thus resembling what observed in the cortex during vibrotactile stimuli in humans and object recognition in monkeys. Memory juggling and competition emerge already by loading only two items. However more items can be stored in WM by considering neural architectures composed of multiple excitatory populations and a common inhibitory pool. Memory capacity depends strongly on the presentation rate of the items and it maximizes for an optimal frequency range. In particular we provide an analytic expression for the maximal memory capacity. Furthermore, the mean membrane potential turns out to be a suitable proxy to measure the memory load, analogously to event driven potentials in experiments on humans. Finally we show that the $\gamma$ power increases with the number of loaded items, as reported in many experiments, while $\theta$ and $\beta$ power reveal non monotonic behaviours.

q-bio.NC↗

arXiv2020

Emergent excitability in adaptive networks of non-excitable units

Population bursts in a large ensemble of coupled elements result from the interplay between the local excitable properties of the nodes and the global network topology. Here collective excitability and self-sustained bursting oscillations are shown to spontaneously emerge in adaptive networks of globally coupled non-excitable units. The ingredients to observe collective excitability are the coexistence of states with different degree of synchronizaton joined to a global feedback acting, on a slow timescale, against the synchronization (desynchronization) of the oscillators. These regimes are illustrated for two paradigmatic classes of coupled rotators: namely, the Kuramoto model with and without inertia. For the bimodal Kuramoto model we analytically show that the macroscopic evolution originates from the existence of a critical manifold organizing the fast collective dynamics on a slow timescale. Our results provide evidence that adaptation can induce excitability by maintaining a network permanently out-of-equilibrium.

nlin.AO↗

arXiv2020

Theta-nested gamma oscillations in next generation neural mass models

Theta-nested gamma oscillations have been reported in many areas of the brain and are believed to represent a fundamental mechanism to transfer information across spatial and temporal scales. In a series of recent experiments in vitro it has been possible to replicate with an optogenetic theta frequency stimulation several features of cross-frequency coupling among theta and gamma rhythms observed in behaving animals. In order to reproduce the main findings of these experiments we have considered a new class of neural mass models able to reproduce exactly the macroscopic dynamics of spiking neural networks. In this framework, we have examined two set-ups able to support collective gamma oscillations: the pyramidal interneuronal network gamma (PING) and the interneuronal network gamma (ING). In both set-ups we observe the emergence of theta-nested gamma oscillations by driving the system with a sinusoidal theta-forcing in proximity of a Hopf bifurcation. These mixed rhythms display always phase amplitude coupling. However 2 different types of nested oscillations can be identified: one characterized by a perfect phase locking between theta and gamma rhythms, corresponding to an overall periodic behaviour; another one where the locking is imperfect and the dynamics is quasi-periodic or even chaotic. From our analysis it emerges that the locked states are more frequent in the ING set-up. In agreement with the experiments, we find theta-nested gamma oscillations for forcing frequencies in the range [1:10] Hz, whose amplitudes grow proportionally to the forcing one and which are clearly modulated by the theta phase. At variance with experimental findings, the gamma-power peak does not shift to higher frequencies by increasing the theta frequency. This effect can be obtained, in or model, only by incrementing, at the same time, also the noise or the forcing amplitude.

q-bio.NC↗