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Tom Tetzlaff

Publications and source records attributed to Tom Tetzlaff.

At least 19 recordsLinked to original sources

Learning sequence timing and control of replay speed in networks of spiking neurons

Processing sequential inputs is a fundamental brain function, underlying tasks such as sensory perception, language, and motor control. A challenge in sequence processing is to represent not only the order of events, but also their precise timing. While existing computational models can learn sequential structure, many lack biologically plausible mechanisms to encode element-specific timing and to flexibly control the speed of sequence replay. The spiking Temporal Memory (sTM) model, a biologically inspired network model, provides a framework for key aspects of sequence processing. In the sTM model, each sequence element is represented by a small set of neurons firing synchronously, where the set of active neurons encodes the element's identity in its sequential context. In its original version, however, the sTM model learns the order but not the timing of sequence elements. Further, it remains an open question in neuroscience how the speed of sequence replay can be flexibly modulated. We propose a mechanism where the duration of sequence elements is represented by a sequential activation of element specific neuronal populations, enabling the model to encode sequences across a wide range of timescales. This provides a biologically plausible basis for learning and replaying complex temporal patterns. Additionally, we show that oscillatory background inputs can serve as a clock signal and provide a robust and flexible mechanism for controlling the speed of sequence replay. Our findings suggest that elapsed time is encoded by unique and sparse spatiotemporal patterns of neural activity, and that the speed of sequence replay during wakefulness and sleep is correlated to the characteristics of global oscillatory activity observed in EEG or LFP recordings.

q-bio.NC

Constructive community race: full-density spiking neural network model drives neuromorphic computing

The local circuitry of the mammalian brain is a focus of the search for generic computational principles because it is largely conserved across species and modalities. In 2014 a model was proposed representing all neurons and synapses of the stereotypical cortical microcircuit below $1\,\text{mm}^2$ of brain surface. The model reproduces fundamental features of brain activity but its impact remained limited because of its computational demands. For theory and simulation, however, the model was a breakthrough because it removes uncertainties of downscaling, and larger models are less densely connected. This sparked a race in the neuromorphic computing community and the model became a de facto standard benchmark. Within a few years real-time performance was reached and surpassed at significantly reduced energy consumption. We review how the computational challenge was tackled by different simulation technologies and derive guidelines for the next generation of benchmarks and other domains of science.

cs.PF

Metadata practices for simulation workflows

Computer simulations are an essential pillar of knowledge generation in science. Exploring, understanding, reproducing, and sharing the results of simulations relies on tracking and organizing the metadata describing the numerical experiments. The models used to understand real-world systems, and the computational machinery required to simulate them, are typically complex, and produce large amounts of heterogeneous metadata. Here, we present general practices for acquiring and handling metadata that are agnostic to software and hardware, and highly flexible for the user. These consist of two steps: 1) recording and storing raw metadata, and 2) selecting and structuring metadata. As a proof of concept, we develop the Archivist, a Python tool to help with the second step, and use it to apply our practices to distinct high-performance computing use cases from neuroscience and hydrology. Our practices and the Archivist can readily be applied to existing workflows without the need for substantial restructuring. They support sustainable numerical workflows, fostering replicability, reproducibility, data exploration, and data sharing in simulation-based research.

cs.IR

Phenomenological modeling of diverse and heterogeneous synaptic dynamics at natural density

This chapter sheds light on the synaptic organization of the brain from the perspective of computational neuroscience. It provides an introductory overview on how to account for empirical data in mathematical models, implement such models in software, and perform simulations reflecting experiments. This path is demonstrated with respect to four key aspects of synaptic signaling: the connectivity of brain networks, synaptic transmission, synaptic plasticity, and the heterogeneity across synapses. Each step and aspect of the modeling and simulation workflow comes with its own challenges and pitfalls, which are highlighted and addressed.

q-bio.NC

Sequence learning in a spiking neuronal network with memristive synapses

Brain-inspired computing proposes a set of algorithmic principles that hold promise for advancing artificial intelligence. They endow systems with self learning capabilities, efficient energy usage, and high storage capacity. A core concept that lies at the heart of brain computation is sequence learning and prediction. This form of computation is essential for almost all our daily tasks such as movement generation, perception, and language. Understanding how the brain performs such a computation is not only important to advance neuroscience but also to pave the way to new technological brain-inspired applications. A previously developed spiking neural network implementation of sequence prediction and recall learns complex, high-order sequences in an unsupervised manner by local, biologically inspired plasticity rules. An emerging type of hardware that holds promise for efficiently running this type of algorithm is neuromorphic hardware. It emulates the way the brain processes information and maps neurons and synapses directly into a physical substrate. Memristive devices have been identified as potential synaptic elements in neuromorphic hardware. In particular, redox-induced resistive random access memories (ReRAM) devices stand out at many aspects. They permit scalability, are energy efficient and fast, and can implement biological plasticity rules. In this work, we study the feasibility of using ReRAM devices as a replacement of the biological synapses in the sequence learning model. We implement and simulate the model including the ReRAM plasticity using the neural simulator NEST. We investigate the effect of different device properties on the performance characteristics of the sequence learning model, and demonstrate resilience with respect to different on-off ratios, conductance resolutions, device variability, and synaptic failure.

cs.NE

Coherent noise enables probabilistic sequence replay in spiking neuronal networks

Animals rely on different decision strategies when faced with ambiguous or uncertain cues. Depending on the context, decisions may be biased towards events that were most frequently experienced in the past, or be more explorative. A particular type of decision making central to cognition is sequential memory recall in response to ambiguous cues. A previously developed spiking neuronal network implementation of sequence prediction and recall learns complex, high-order sequences in an unsupervised manner by local, biologically inspired plasticity rules. In response to an ambiguous cue, the model deterministically recalls the sequence shown most frequently during training. Here, we present an extension of the model enabling a range of different decision strategies. In this model, explorative behavior is generated by supplying neurons with noise. As the model relies on population encoding, uncorrelated noise averages out, and the recall dynamics remain effectively deterministic. In the presence of locally correlated noise, the averaging effect is avoided without impairing the model performance, and without the need for large noise amplitudes. We investigate two forms of correlated noise occurring in nature: shared synaptic background inputs, and random locking of the stimulus to spatiotemporal oscillations in the network activity. Depending on the noise characteristics, the network adopts various replay strategies. This study thereby provides potential mechanisms explaining how the statistics of learned sequences affect decision making, and how decision strategies can be adjusted after learning.

q-bio.NC

A Modular Workflow for Performance Benchmarking of Neuronal Network Simulations

Modern computational neuroscience strives to develop complex network models to explain dynamics and function of brains in health and disease. This process goes hand in hand with advancements in the theory of neuronal networks and increasing availability of detailed anatomical data on brain connectivity. Large-scale models that study interactions between multiple brain areas with intricate connectivity and investigate phenomena on long time scales such as system-level learning require progress in simulation speed. The corresponding development of state-of-the-art simulation engines relies on information provided by benchmark simulations which assess the time-to-solution for scientifically relevant, complementary network models using various combinations of hardware and software revisions. However, maintaining comparability of benchmark results is difficult due to a lack of standardized specifications for measuring the scaling performance of simulators on high-performance computing (HPC) systems. Motivated by the challenging complexity of benchmarking, we define a generic workflow that decomposes the endeavor into unique segments consisting of separate modules. As a reference implementation for the conceptual workflow, we develop beNNch: an open-source software framework for the configuration, execution, and analysis of benchmarks for neuronal network simulations. The framework records benchmarking data and metadata in a unified way to foster reproducibility. For illustration, we measure the performance of various versions of the NEST simulator across network models with different levels of complexity on a contemporary HPC system, demonstrating how performance bottlenecks can be identified, ultimately guiding the development toward more efficient simulation technology.

q-bio.NC

Sequence learning, prediction, and replay in networks of spiking neurons

Sequence learning, prediction and replay have been proposed to constitute the universal computations performed by the neocortex. The Hierarchical Temporal Memory (HTM) algorithm realizes these forms of computation. It learns sequences in an unsupervised and continuous manner using local learning rules, permits a context specific prediction of future sequence elements, and generates mismatch signals in case the predictions are not met. While the HTM algorithm accounts for a number of biological features such as topographic receptive fields, nonlinear dendritic processing, and sparse connectivity, it is based on abstract discrete-time neuron and synapse dynamics, as well as on plasticity mechanisms that can only partly be related to known biological mechanisms. Here, we devise a continuous-time implementation of the temporal-memory (TM) component of the HTM algorithm, which is based on a recurrent network of spiking neurons with biophysically interpretable variables and parameters. The model learns high-order sequences by means of a structural Hebbian synaptic plasticity mechanism supplemented with a rate-based homeostatic control. In combination with nonlinear dendritic input integration and local inhibitory feedback, this type of plasticity leads to the dynamic self-organization of narrow sequence-specific subnetworks. These subnetworks provide the substrate for a faithful propagation of sparse, synchronous activity, and, thereby, for a robust, context specific prediction of future sequence elements as well as for the autonomous replay of previously learned sequences. By strengthening the link to biology, our implementation facilitates the evaluation of the TM hypothesis based on electrophysiological and behavioral data.

q-bio.NC

Prominent characteristics of recurrent neuronal networks are robust against low synaptic weight resolution

The representation of the natural-density, heterogeneous connectivity of neuronal network models at relevant spatial scales remains a challenge for Computational Neuroscience and Neuromorphic Computing. In particular, the memory demands imposed by the vast number of synapses in brain-scale network simulations constitutes a major obstacle. Limiting the number resolution of synaptic weights appears to be a natural strategy to reduce memory and compute load. In this study, we investigate the effects of a limited synaptic-weight resolution on the dynamics of recurrent spiking neuronal networks resembling local cortical circuits, and develop strategies for minimizing deviations from the dynamics of networks with high-resolution synaptic weights. We mimic the effect of a limited synaptic weight resolution by replacing normally distributed synaptic weights by weights drawn from a discrete distribution, and compare the resulting statistics characterizing firing rates, spike-train irregularity, and correlation coefficients with the reference solution. We show that a naive discretization of synaptic weights generally leads to a distortion of the spike-train statistics. Only if the weights are discretized such that the mean and the variance of the total synaptic input currents are preserved, the firing statistics remains unaffected for the types of networks considered in this study. For networks with sufficiently heterogeneous in-degrees, the firing statistics can be preserved even if all synaptic weights are replaced by the mean of the weight distribution. We conclude that even for simple networks with non-plastic neurons and synapses, a discretization of synaptic weights can lead to substantial deviations in the firing statistics, unless the discretization is performed with care and guided by a rigorous validation process.

q-bio.NC

Firing rate homeostasis counteracts changes in stability of recurrent neural networks caused by synapse loss in Alzheimer's disease

The impairment of cognitive function in Alzheimer's is clearly correlated to synapse loss. However, the mechanisms underlying this correlation are only poorly understood. Here, we investigate how the loss of excitatory synapses in sparsely connected random networks of spiking excitatory and inhibitory neurons alters their dynamical characteristics. Beyond the effects on the network's activity statistics, we find that the loss of excitatory synapses on excitatory neurons shifts the network dynamic towards the stable regime. The decrease in sensitivity to small perturbations to time varying input can be considered as an indication of a reduction of computational capacity. A full recovery of the network performance can be achieved by firing rate homeostasis, here implemented by an up-scaling of the remaining excitatory-excitatory synapses. By analysing the stability of the linearized network dynamics, we explain how homeostasis can simultaneously maintain the network's firing rate and sensitivity to small perturbations.

q-bio.NC

Conditions for wave trains in spiking neural networks

Spatiotemporal patterns such as traveling waves are frequently observed in recordings of neural activity. The mechanisms underlying the generation of such patterns are largely unknown. Previous studies have investigated the existence and uniqueness of different types of waves or bumps of activity using neural-field models, phenomenological coarse-grained descriptions of neural-network dynamics. But it remains unclear how these insights can be transferred to more biologically realistic networks of spiking neurons, where individual neurons fire irregularly. Here, we employ mean-field theory to reduce a microscopic model of leaky integrate-and-fire (LIF) neurons with distance-dependent connectivity to an effective neural-field model. In contrast to existing phenomenological descriptions, the dynamics in this neural-field model depends on the mean and the variance in the synaptic input, both determining the amplitude and the temporal structure of the resulting effective coupling kernel. For the neural-field model we employ liner stability analysis to derive conditions for the existence of spatial and temporal oscillations and wave trains, that is, temporally and spatially periodic traveling waves. We first prove that wave trains cannot occur in a single homogeneous population of neurons, irrespective of the form of distance dependence of the connection probability. Compatible with the architecture of cortical neural networks, wave trains emerge in two-population networks of excitatory and inhibitory neurons as a combination of delay-induced temporal oscillations and spatial oscillations due to distance-dependent connectivity profiles. Finally, we demonstrate quantitative agreement between predictions of the analytically tractable neural-field model and numerical simulations of both networks of nonlinear rate-based units and networks of LIF neurons.

q-bio.NC

Deterministic networks for probabilistic computing

Neural-network models of high-level brain functions such as memory recall and reasoning often rely on the presence of stochasticity. The majority of these models assumes that each neuron in the functional network is equipped with its own private source of randomness, often in the form of uncorrelated external noise. However, both in vivo and in silico, the number of noise sources is limited due to space and bandwidth constraints. Hence, neurons in large networks usually need to share noise sources. Here, we show that the resulting shared-noise correlations can significantly impair the performance of stochastic network models. We demonstrate that this problem can be overcome by using deterministic recurrent neural networks as sources of uncorrelated noise, exploiting the decorrelating effect of inhibitory feedback. Consequently, even a single recurrent network of a few hundred neurons can serve as a natural noise source for large ensembles of functional networks, each comprising thousands of units. We successfully apply the proposed framework to a diverse set of binary-unit networks with different dimensionalities and entropies, as well as to a network reproducing handwritten digits with distinct predefined frequencies. Finally, we show that the same design transfers to functional networks of spiking neurons.

q-bio.NC

The effect of heterogeneity on decorrelation mechanisms in spiking neural networks: a neuromorphic-hardware study

High-level brain function such as memory, classification or reasoning can be realized by means of recurrent networks of simplified model neurons. Analog neuromorphic hardware constitutes a fast and energy efficient substrate for the implementation of such neural computing architectures in technical applications and neuroscientific research. The functional performance of neural networks is often critically dependent on the level of correlations in the neural activity. In finite networks, correlations are typically inevitable due to shared presynaptic input. Recent theoretical studies have shown that inhibitory feedback, abundant in biological neural networks, can actively suppress these shared-input correlations and thereby enable neurons to fire nearly independently. For networks of spiking neurons, the decorrelating effect of inhibitory feedback has so far been explicitly demonstrated only for homogeneous networks of neurons with linear sub-threshold dynamics. Theory, however, suggests that the effect is a general phenomenon, present in any system with sufficient inhibitory feedback, irrespective of the details of the network structure or the neuronal and synaptic properties. Here, we investigate the effect of network heterogeneity on correlations in sparse, random networks of inhibitory neurons with non-linear, conductance-based synapses. Emulations of these networks on the analog neuromorphic hardware system Spikey allow us to test the efficiency of decorrelation by inhibitory feedback in the presence of hardware-specific heterogeneities. The configurability of the hardware substrate enables us to modulate the extent of heterogeneity in a systematic manner. We selectively study the effects of shared input and recurrent connections on correlations in membrane potentials and spike trains. Our results confirm ...

q-bio.NC

Hybrid scheme for modeling local field potentials from point-neuron networks

Due to rapid advances in multielectrode recording technology, the local field potential (LFP) has again become a popular measure of neuronal activity in both basic research and clinical applications. Proper understanding of the LFP requires detailed mathematical modeling incorporating the anatomical and electrophysiological features of neurons near the recording electrode, as well as synaptic inputs from the entire network. Here we propose a hybrid modeling scheme combining the efficiency of commonly used simplified point-neuron network models with the biophysical principles underlying LFP generation by real neurons. The scheme can be used with an arbitrary number of point-neuron network populations. The LFP predictions rely on populations of network-equivalent, anatomically reconstructed multicompartment neuron models with layer-specific synaptic connectivity. The present scheme allows for a full separation of the network dynamics simulation and LFP generation. For illustration, we apply the scheme to a full-scale cortical network model for a $\sim$1 mm$^2$ patch of primary visual cortex and predict laminar LFPs for different network states, assess the relative LFP contribution from different laminar populations, and investigate the role of synaptic input correlations and neuron density on the LFP. The generic nature of the hybrid scheme and its publicly available implementation in \texttt{hybridLFPy} form the basis for LFP predictions from other point-neuron network models, as well as extensions of the current application to larger circuitry and additional biological detail.

q-bio.NC

On 1/f^alpha power laws originating from linear neuronal cable theory: power spectral densities of the soma potential, transmembrane current and single-neuron contribution to the EEG

Power laws, that is, power spectral densities (PSDs) exhibiting 1/f^alpha behavior for large frequencies f, have commonly been observed in neural recordings. Power laws in noise spectra have not only been observed in microscopic recordings of neural membrane potentials and membrane currents, but also in macroscopic EEG (electroencephalographic) recordings. While complex network behavior has been suggested to be at the root of this phenomenon, we here demonstrate a possible origin of such power laws in the biophysical properties of single neurons described by the standard cable equation. Taking advantage of the analytical tractability of the so called ball and stick neuron model, we derive general expressions for the PSD transfer functions for a set of measures of neuronal activity: the soma membrane current, the current-dipole moment (corresponding to the single-neuron EEG contribution), and the soma membrane potential. These PSD transfer functions relate the PSDs of the respective measurements to the PSDs of the noisy input currents. With homogeneously distributed input currents across the neuronal membrane we find that all PSD transfer functions express asymptotic high-frequency 1/f^alpha power laws. The corresponding power-law exponents are analytically identified as alpha_inf^I = 1/2 for the soma membrane current, alpha_inf^p = 3/2 for the current-dipole moment, and alpha_inf^V = 2 for the soma membrane potential. These power-law exponents are found for arbitrary combinations of uncorrelated and correlated noisy input current (as long as both the dendrites and the soma receive some uncorrelated input currents). Comparison with available data suggests that the apparent power laws observed in experiments may stem from uncorrelated current sources, presumably intrinsic ion channels, which are homogeneously distributed across the neural membranes and themselves exhibit ...

physics.bio-ph

A unified view on weakly correlated recurrent networks

The diversity of neuron models used in contemporary theoretical neuroscience to investigate specific properties of covariances raises the question how these models relate to each other. In particular it is hard to distinguish between generic properties and peculiarities due to the abstracted model. Here we present a unified view on pairwise covariances in recurrent networks in the irregular regime. We consider the binary neuron model, the leaky integrate-and-fire model, and the Hawkes process. We show that linear approximation maps each of these models to either of two classes of linear rate models, including the Ornstein-Uhlenbeck process as a special case. The classes differ in the location of additive noise in the rate dynamics, which is on the output side for spiking models and on the input side for the binary model. Both classes allow closed form solutions for the covariance. For output noise it separates into an echo term and a term due to correlated input. The unified framework enables us to transfer results between models. For example, we generalize the binary model and the Hawkes process to the presence of conduction delays and simplify derivations for established results. Our approach is applicable to general network structures and suitable for population averages. The derived averages are exact for fixed out-degree network architectures and approximate for fixed in-degree. We demonstrate how taking into account fluctuations in the linearization procedure increases the accuracy of the effective theory and we explain the class dependent differences between covariances in the time and the frequency domain. Finally we show that the oscillatory instability emerging in networks of integrate-and-fire models with delayed inhibitory feedback is a model-invariant feature: the same structure of poles in the complex frequency plane determines the population power spectra.

q-bio.NC

The correlation structure of local cortical networks intrinsically results from recurrent dynamics

The co-occurrence of action potentials of pairs of neurons within short time intervals is known since long. Such synchronous events can appear time-locked to the behavior of an animal and also theoretical considerations argue for a functional role of synchrony. Early theoretical work tried to explain correlated activity by neurons transmitting common fluctuations due to shared inputs. This, however, overestimates correlations. Recently the recurrent connectivity of cortical networks was shown responsible for the observed low baseline correlations. Two different explanations were given: One argues that excitatory and inhibitory population activities closely follow the external inputs to the network, so that their effects on a pair of cells mutually cancel. Another explanation relies on negative recurrent feedback to suppress fluctuations in the population activity, equivalent to small correlations. In a biological neuronal network one expects both, external inputs and recurrence, to affect correlated activity. The present work extends the theoretical framework of correlations to include both contributions and explains their qualitative differences. Moreover the study shows that the arguments of fast tracking and recurrent feedback are not equivalent, only the latter correctly predicts the cell-type specific correlations.

q-bio.NC

Echoes in correlated neural systems

Correlations are employed in modern physics to explain microscopic and macroscopic phenomena, like the fractional quantum Hall effect and the Mott insulator state in high temperature superconductors and ultracold atoms. Simultaneously probed neurons in the intact brain reveal correlations between their activity, an important measure to study information processing in the brain that also influences macroscopic signals of neural activity, like the electro encephalogram (EEG). Networks of spiking neurons differ from most physical systems: The interaction between elements is directed, time delayed, mediated by short pulses, and each neuron receives events from thousands of neurons. Even the stationary state of the network cannot be described by equilibrium statistical mechanics. Here we develop a quantitative theory of pairwise correlations in finite sized random networks of spiking neurons. We derive explicit analytic expressions for the population averaged cross correlation functions. Our theory explains why the intuitive mean field description fails, how the echo of single action potentials causes an apparent lag of inhibition with respect to excitation, and how the size of the network can be scaled while maintaining its dynamical state. Finally, we derive a new criterion for the emergence of collective oscillations from the spectrum of the time-evolution propagator.

q-bio.NC