arXiv subjects

Tilo Schwalger

Publications and source records attributed to Tilo Schwalger.

12 recordsLinked to original sources

arXiv2024

How random connectivity shapes the fluctuating dynamics of finite-size neural populations

Mesoscopic models of finite-size neuronal populations are crucial to understand the dynamics of neural networks in the brain, especially their fluctuations and response to stimuli. However, current theories to derive such models are based on homogeneous all-to-all (full) connectivity. This assumption neglects the variance in the connectivity of biologically realistic networks with connection probabilities $p<1$ (non-full connectivity). To gain insight into the different fluctuation mechanisms underlying neural variability at the population level, we derive and analyze a stochastic mean-field model for finite-size networks of Poisson neurons with random connectivity (including non-full connectivity), external noise and disordered mean inputs. We treat the quenched disorder of the connectivity by an annealed approximation enabling a doubly stochastic description of synaptic inputs for finite network size. A further reduction leads to a low-dimensional closed system of coupled Langevin equations for the mean and variance of the membrane potentials as well as a variable capturing finite-size fluctuations. Compared to microscopic simulations, the mesoscopic model describes the fluctuations and nonlinearities well and outperforms previous theories that neglected the variance in the connectivity. The joint effect of connectivity disorder and finite network size can be analytically understood by a softening of the effective nonlinearity and the multiplicative character of spiking noise. The mesoscopic theory shows that quenched disorder can stabilize the asynchronous state, and it correctly predicts large quantitative and non-trivial qualitative effects of connection probability on the variance of the population firing rate and its dependence on stimulus strength. In conclusion, our theory elucidates how disordered connectivity shapes nonlinear dynamics and fluctuations of neural populations.

q-bio.NC↗

arXiv2024

A refractory density approach to a multi-scale SEIRS epidemic model

We propose a novel multi-scale modeling framework for infectious disease spreading, borrowing ideas and modeling tools from the so-called Refractory Density (RD) approach. We introduce a microscopic model that describes the probability of infection for a single individual and the evolution of the disease within their body. From the individual-level description, we then present the corresponding population-level model of epidemic spreading on the mesoscopic and macroscopic scale. We conclude with numerical illustrations taking into account either a white Gaussian noise or an escape noise to showcase the potential of our approach in producing both transient and asymptotic complex dynamics as well as finite-size fluctuations consistently across multiple scales. A comparison with the epidemiology of coronaviruses is also given to corroborate the qualitative relevance of our new approach.

q-bio.PE↗

arXiv2022

On a finite-size neuronal population equation

Population equations for infinitely large networks of spiking neurons have a long tradition in theoretical neuroscience. In this work, we analyze a recent generalization of these equations to populations of finite size, which takes the form of a nonlinear stochastic integral equation. We prove that, in the case of leaky integrate-and-fire (LIF) neurons with escape noise and for a slightly simplified version of the model, the equation is well-posed and stable in the sense of Brémaud-Massoulié. The proof combines methods from Markov processes taking values in the space of positive measures and nonlinear Hawkes processes. For applications, we also provide efficient simulation algorithms.

math.PR↗

arXiv2022

Mesoscopic description of hippocampal replay and metastability in spiking neural networks with short-term plasticity

Bottom-up models of functionally relevant patterns of neural activity provide an explicit link between neuronal dynamics and computation. A prime example of functional activity pattern is hippocampal replay, which is critical for memory consolidation. The switchings between replay events and a low-activity state in neural recordings suggests metastable neural circuit dynamics. As metastability has been attributed to noise and/or slow fatigue mechanisms, we propose a concise mesoscopic model which accounts for both. Crucially, our model is bottom-up: it is analytically derived from the dynamics of finite-size networks of Linear-Nonlinear Poisson neurons with short-term synaptic depression. As such, noise is explicitly linked to spike noise and network size, and fatigue is explicitly linked to synaptic dynamics. To derive the mesosocpic model, we first consider a homogeneous spiking neural network and follow the temporal coarse-graining approach of Gillespie ("chemical Langevin equation"), which can be naturally interpreted as a stochastic neural mass model. The Langevin equation is computationally inexpensive to simulate and enables a thorough study of metastable dynamics in classical setups (population spikes and Up-Down states dynamics) by means of phase-plane analysis. This stochastic neural mass model is the basic component of our mesoscopic model for replay. We show that our model faithfully captures the stochastic nature of individual replayed trajectories. Moreover, compared to the deterministic Romani-Tsodyks model of place cell dynamics, it exhibits a higher level of variability in terms of content, direction and timing of replay events, compatible with biological evidence and could be functionally desirable. This variability is the product of a new dynamical regime where metastability emerges from a complex interplay between finite-size fluctuations and local fatigue.

q-bio.NC↗

arXiv2021

Mapping Input Noise to Escape Noise in Integrate-and-fire neurons: A Level-Crossing Approach

Noise in spiking neurons is commonly modeled by a noisy input current or by generating output spikes stochastically with a voltage-dependent hazard rate ("escape noise"). While input noise lends itself to modeling biophysical noise processes, the phenomenological escape noise is mathematically more tractable. Using the level-crossing theory for differentiable Gaussian processes, we derive an approximate mapping between colored input noise and escape noise in leaky integrate-and-fire neurons. This mapping requires the first-passage-time (FPT) density of an overdamped Brownian particle driven by colored noise with respect to an arbitrarily moving boundary. Starting from the Wiener-Rice series for the FPT density, we apply the second-order decoupling approximation of Stratonovich to the case of moving boundaries and derive a simplified hazard-rate representation that is local in time and numerically efficient. This simplification requires the calculation of the non-stationary auto-correlation function of the level-crossing process: For exponentially correlated input noise (Ornstein-Uhlenbeck process), we obtain an exact formula for the zero-lag auto-correlation as a function of noise parameters, mean membrane potential and its speed, as well as an exponential approximation of the full auto-correlation function. The theory well predicts the FPT and interspike interval densities as well as the population activities obtained from simulations with time-dependent stimulus or boundary. The agreement with simulations is strongly enhanced compared to a first-order decoupling approximation that neglects correlations between level crossings. The second-order approximation also improves upon a previously proposed theory in the subthreshold regime. Depending on a simplicity-accuracy trade-off, all considered approximations represent useful mappings from colored input noise to escape noise.

q-bio.NC↗

arXiv2020

Low-dimensional firing-rate dynamics for populations of renewal-type spiking neurons

The macroscopic dynamics of large populations of neurons can be mathematically analyzed using low-dimensional firing-rate or neural-mass models. However, these models fail to capture spike synchronization effects of stochastic spiking neurons such as the non-stationary population response to rapidly changing stimuli. Here, we derive low-dimensional firing-rate models for homogeneous populations of general renewal-type neurons, including integrate-and-fire models driven by white noise. Renewal models account for neuronal refractoriness and spike synchronization dynamics. The derivation is based on an eigenmode expansion of the associated refractory density equation, which generalizes previous spectral methods for Fokker-Planck equations to arbitrary renewal models. We find a simple relation between the eigenvalues, which determine the characteristic time scales of the firing rate dynamics, and the Laplace transform of the interspike interval density or the survival function of the renewal process. Analytical expressions for the Laplace transforms are readily available for many renewal models including the leaky integrate-and-fire model. Retaining only the first eigenmode yields already an adequate low-dimensional approximation of the firing-rate dynamics that captures spike synchronization effects and fast transient dynamics at stimulus onset. We explicitly demonstrate the validity of our model for a large homogeneous population of Poisson neurons with absolute refractoriness, and other renewal models that admit an explicit analytical calculation of the eigenvalues. The here presented eigenmode expansion provides a systematic framework for novel firing-rate models in computational neuroscience based on spiking neuron dynamics with refractoriness.

q-bio.NC↗

arXiv2019

Mind the Last Spike -- Firing Rate Models for Mesoscopic Populations of Spiking Neurons

The dominant modeling framework for understanding cortical computations are heuristic firing rate models. Despite their success, these models fall short to capture spike synchronization effects, to link to biophysical parameters and to describe finite-size fluctuations. In this opinion article, we propose that the refractory density method (RDM), also known as age-structured population dynamics or quasi-renewal theory, yields a powerful theoretical framework to build rate-based models for mesoscopic neural populations from realistic neuron dynamics at the microscopic level. We review recent advances achieved by the RDM to obtain efficient population density equations for networks of generalized integrate-and-fire (GIF) neurons -- a class of neuron models that has been successfully fitted to various cell types. The theory not only predicts the nonstationary dynamics of large populations of neurons but also permits an extension to finite-size populations and a systematic reduction to low-dimensional rate dynamics. The new types of rate models will allow a re-examination of models of cortical computations under biological constraints.

q-bio.NC↗

arXiv2019

How single neuron properties shape chaotic dynamics and signal transmission in random neural networks

While most models of randomly connected networks assume nodes with simple dynamics, nodes in realistic highly connected networks, such as neurons in the brain, exhibit intrinsic dynamics over multiple timescales. We analyze how the dynamical properties of nodes (such as single neurons) and recurrent connections interact to shape the effective dynamics in large randomly connected networks. A novel dynamical mean-field theory for strongly connected networks of multi-dimensional rate units shows that the power spectrum of the network activity in the chaotic phase emerges from a nonlinear sharpening of the frequency response function of single units. For the case of two-dimensional rate units with strong adaptation, we find that the network exhibits a state of "resonant chaos", characterized by robust, narrow-band stochastic oscillations. The coherence of stochastic oscillations is maximal at the onset of chaos and their correlation time scales with the adaptation timescale of single units. Surprisingly, the resonance frequency can be predicted from the properties of isolated units, even in the presence of heterogeneity in the adaptation parameters. In the presence of these internally-generated chaotic fluctuations, the transmission of weak, low-frequency signals is strongly enhanced by adaptation, whereas signal transmission is not influenced by adaptation in the non-chaotic regime. Our theoretical framework can be applied to other mechanisms at the level of single nodes, such as synaptic filtering, refractoriness or spike synchronization. These results advance our understanding of the interaction between the dynamics of single units and recurrent connectivity, which is a fundamental step toward the description of biologically realistic network models in the brain, or, more generally, networks of other physical or man-made complex dynamical units.

cond-mat.dis-nn↗

arXiv2018

Mesoscopic population equations for spiking neural networks with synaptic short-term plasticity

Coarse-graining microscopic models of biological neural networks to obtain mesoscopic models of neural activities is an essential step towards multi-scale models of the brain. Here, we extend a recent theory for mesoscopic population dynamics with static synapses to the case of dynamic synapses exhibiting short-term plasticity (STP). Under the assumption that spike arrivals at synapses have Poisson statistics, we derive analytically stochastic mean-field dynamics for the effective synaptic coupling between finite-size populations undergoing Tsodyks-Markram STP. The novel mean-field equations account for both finite number of synapses and correlations between the neurotransmitter release probability and the fraction of available synaptic resources. Comparisons with Monte Carlo simulations of the microscopic model show that in both feedforward and recurrent networks the mesoscopic mean-field model accurately reproduces stochastic realizations of the total synaptic input into a postsynaptic neuron and accounts for stochastic switches between Up and Down states as well as for population spikes. The extended mesoscopic population theory of spiking neural networks with STP may be useful for a systematic reduction of detailed biophysical models of cortical microcircuits to efficient and mathematically tractable mean-field models.

q-bio.NC↗

arXiv2017

Patterns of interval correlations in neural oscillators with adaptation

Neural firing is often subject to negative feedback by adaptation currents. These currents can induce strong correlations among the time intervals between spikes. Here we study analytically the interval correlations of a broad class of noisy neural oscillators with spike-triggered adaptation of arbitrary strength and time scale. Our weak-noise theory provides a general relation between the correlations and the phase-response curve (PRC) of the oscillator, proves anti-correlations between neighboring intervals for adapting neurons with type I PRC and identifies a single order parameter that determines the qualitative pattern of correlations. Monotonically decaying or oscillating correlation structures can be related to qualitatively different voltage traces after spiking, which can be explained by the phase plane geometry. At high firing rates, the long-term variability of the spike train associated with the cumulative interval correlations becomes small, independent of model details. Our results are verified by comparison with stochastic simulations of the exponential, leaky, and generalized integrate-and-fire models with adaptation.

q-bio.NC↗

arXiv2017

Towards a theory of cortical columns: From spiking neurons to interacting neural populations of finite size

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 effects in single-neuron dynamics such as refractoriness and adaptation interact with finite-size fluctuations on the population level. Efficient integration of the stochastic mesoscopic equations reproduces the statistical behavior of the population activities obtained from microscopic simulations of a full spiking neural network model. The theory describes nonlinear emergent dynamics like finite-size-induced stochastic transitions in multistable networks and synchronization in balanced networks of excitatory and inhibitory neurons. The mesoscopic equations are employed to rapidly simulate a model of a local cortical microcircuit consisting of eight neuron types. Our theory establishes a general framework for modeling finite-size neural population dynamics based on single cell and synapse parameters and offers an efficient approach to analyzing cortical circuits and computations.

q-bio.NC↗

arXiv2015

Fluctuations and information filtering in coupled populations of spiking neurons with adaptation

Finite-sized populations of spiking elements are fundamental to brain function, but also used in many areas of physics. Here we present a theory of the dynamics of finite-sized populations of spiking units, based on a quasi-renewal description of neurons with adaptation. We derive an integral equation with colored noise that governs the stochastic dynamics of the population activity in response to time-dependent stimulation and calculate the spectral density in the asynchronous state. We show that systems of coupled populations with adaptation can generate a frequency band in which sensory information is preferentially encoded. The theory is applicable to fully as well as randomly connected networks, and to leaky integrate-and-fire as well as to generalized spiking neurons with adaptation on multiple time scales.

q-bio.NC↗