arXiv subjects

Wulfram Gerstner

Publications and source records attributed to Wulfram Gerstner.

At least 55 records · Page 3Linked to original sources

arXiv2017

Efficient low-dimensional approximation of continuous attractor networks

Continuous "bump" attractors are an established model of cortical working memory for continuous variables and can be implemented using various neuron and network models. Here, we develop a generalizable approach for the approximation of bump states of continuous attractor networks implemented in networks of both rate-based and spiking neurons. The method relies on a low-dimensional parametrization of the spatial shape of firing rates, allowing to apply efficient numerical optimization methods. Using our theory, we can establish a mapping between network structure and attractor properties that allows the prediction of the effects of network parameters on the steady state firing rate profile and the existence of bumps, and vice-versa, to fine-tune a network to produce bumps of a given shape.

q-bio.NC↗

arXiv2017

Predicting non-linear dynamics by stable local learning in a recurrent spiking neural network

Brains need to predict how the body reacts to motor commands. It is an open question how networks of spiking neurons can learn to reproduce the non-linear body dynamics caused by motor commands, using local, online and stable learning rules. Here, we present a supervised learning scheme for the feedforward and recurrent connections in a network of heterogeneous spiking neurons. The error in the output is fed back through fixed random connections with a negative gain, causing the network to follow the desired dynamics, while an online and local rule changes the weights. The rule for Feedback-based Online Local Learning Of Weights (FOLLOW) is local in the sense that weight changes depend on the presynaptic activity and the error signal projected onto the postsynaptic neuron. We provide examples of learning linear, non-linear and chaotic dynamics, as well as the dynamics of a two-link arm. Using the Lyapunov method, and under reasonable assumptions and approximations, we show that FOLLOW learning is stable uniformly, with the error going to zero asymptotically.

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.

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arXiv2017

Automated point-neuron simplification of data-driven microcircuit models

A method is presented for the reduction of morphologically detailed microcircuit models to a point-neuron representation without human intervention. The simplification occurs in a modular workflow, in the neighborhood of a user specified network activity state for the reference model, the "operating point". First, synapses are moved to the soma, correcting for dendritic filtering by low-pass filtering the delivered synaptic current. Filter parameters are computed numerically and independently for inhibitory and excitatory input using a Green's function approach. Next, point-neuron models for each neuron in the microcircuit are fit to their respective morphologically detailed counterparts. Here, generalized integrate-and-fire point neuron models are used, leveraging a recently published fitting toolbox. The fits are constrained by currents and voltages computed in the morphologically detailed partner neurons with soma corrected synapses at three depolarizations about the user specified operating point. The result is a simplified circuit which is well constrained by the reference circuit, and can be continuously updated as the latter iteratively integrates new data. The modularity of the approach makes it applicable also for other point-neuron and synapse models. The approach is demonstrated on a recently reported reconstruction of a neocortical microcircuit around an in vivo-like working point. The resulting simplified network model is benchmarked to the reference morphologically detailed microcircuit model for a range of simulated network protocols. The simplified network is found to be slightly more sub-critical than the reference, with otherwise good agreement for both quantitative and qualitative validations.

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arXiv2017

Balancing New Against Old Information: The Role of Surprise in Learning

Surprise describes a range of phenomena from unexpected events to behavioral responses. We propose a measure of surprise and use it for surprise-driven learning. Our surprise measure takes into account data likelihood as well as the degree of commitment to a belief via the entropy of the belief distribution. We find that surprise-minimizing learning dynamically adjusts the balance between new and old information without the need of knowledge about the temporal statistics of the environment. We apply our framework to a dynamic decision-making task and a maze exploration task. Our surprise minimizing framework is suitable for learning in complex environments, even if the environment undergoes gradual or sudden changes and could eventually provide a framework to study the behavior of humans and animals encountering surprising events.

stat.ML↗

arXiv2016

Towards deep learning with spiking neurons in energy based models with contrastive Hebbian plasticity

In machine learning, error back-propagation in multi-layer neural networks (deep learning) has been impressively successful in supervised and reinforcement learning tasks. As a model for learning in the brain, however, deep learning has long been regarded as implausible, since it relies in its basic form on a non-local plasticity rule. To overcome this problem, energy-based models with local contrastive Hebbian learning were proposed and tested on a classification task with networks of rate neurons. We extended this work by implementing and testing such a model with networks of leaky integrate-and-fire neurons. Preliminary results indicate that it is possible to learn a non-linear regression task with hidden layers, spiking neurons and a local synaptic plasticity rule.

cs.LG↗

arXiv2016

Multi-contact synapses for stable networks: a spike-timing dependent model of dendritic spine plasticity and turnover

Excitatory synaptic connections in the adult neocortex consist of multiple synaptic contacts, almost exclusively formed on dendritic spines. Changes of dendritic spine shape and volume, a correlate of synaptic strength, can be tracked in vivo for weeks. Here, we present a combined model of spike-timing dependent dendritic spine plasticity and turnover that explains the steady state multi-contact configuration of synapses in adult neocortical networks. In this model, many presynaptic neurons compete to make strong synaptic connections onto postsynaptic neurons, while the synaptic contacts comprising each connection cooperate via postsynaptic firing. We demonstrate that the model is consistent with experimentally observed long-term dendritic spine dynamics under steady-state and lesion induced conditions, and show that cooperation of multiple synaptic contacts is crucial for stable, long-term synaptic memories. In simulations of a simplified network of barrel cortex, our plasticity rule reproduces whisker-trimming induced rewiring of thalamo-cortical and recurrent synaptic connectivity on realistic time scales.

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arXiv2016

Algorithmic Composition of Melodies with Deep Recurrent Neural Networks

A big challenge in algorithmic composition is to devise a model that is both easily trainable and able to reproduce the long-range temporal dependencies typical of music. Here we investigate how artificial neural networks can be trained on a large corpus of melodies and turned into automated music composers able to generate new melodies coherent with the style they have been trained on. We employ gated recurrent unit networks that have been shown to be particularly efficient in learning complex sequential activations with arbitrary long time lags. Our model processes rhythm and melody in parallel while modeling the relation between these two features. Using such an approach, we were able to generate interesting complete melodies or suggest possible continuations of a melody fragment that is coherent with the characteristics of the fragment itself.

stat.ML↗

arXiv2016

Nonlinear Hebbian learning as a unifying principle in receptive field formation

The development of sensory receptive fields has been modeled in the past by a variety of models including normative models such as sparse coding or independent component analysis and bottom-up models such as spike-timing dependent plasticity or the Bienenstock-Cooper-Munro model of synaptic plasticity. Here we show that the above variety of approaches can all be unified into a single common principle, namely Nonlinear Hebbian Learning. When Nonlinear Hebbian Learning is applied to natural images, receptive field shapes were strongly constrained by the input statistics and preprocessing, but exhibited only modest variation across different choices of nonlinearities in neuron models or synaptic plasticity rules. Neither overcompleteness nor sparse network activity are necessary for the development of localized receptive fields. The analysis of alternative sensory modalities such as auditory models or V2 development lead to the same conclusions. In all examples, receptive fields can be predicted a priori by reformulating an abstract model as nonlinear Hebbian learning. Thus nonlinear Hebbian learning and natural statistics can account for many aspects of receptive field formation across models and sensory modalities.

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↗

arXiv2013

Spike timing prediction with active dendrites

A complete single-neuron model must correctly reproduce the firing of spikes and bursts. We present a study of a simplified model of deep pyramidal cells of the cortex with active dendrites. We hypothesized that we can model the soma and its apical tuft with only two compartments, without significant loss in the accuracy of spike-timing predictions. The model is based on experimentally measurable impulse-response functions, which transfer the effect of current injected in one compartment to current reaching the other. Each compartment was modeled with a pair of non-linear differential equations and a small number of parameters that approximate the Hodgkin-and-Huxley equations. The predictive power of this model was tested on electrophysiological experiments where noisy current was injected in both the soma and the apical dendrite simultaneously. We conclude that a simple two-compartment model can predict spike times of pyramidal cells stimulated in the soma and dendrites simultaneously. Our results support that regenerating activity in the dendritic tuft is required to properly account for the dynamics of layer 5 pyramidal cells under in-vivo-like conditions.

q-bio.NC↗

arXiv2013

Reward-based learning under hardware constraints - Using a RISC processor embedded in a neuromorphic substrate

In this study, we propose and analyze in simulations a new, highly flexible method of implementing synaptic plasticity in a wafer-scale, accelerated neuromorphic hardware system. The study focuses on globally modulated STDP, as a special use-case of this method. Flexibility is achieved by embedding a general-purpose processor dedicated to plasticity into the wafer. To evaluate the suitability of the proposed system, we use a reward modulated STDP rule in a spike train learning task. A single layer of neurons is trained to fire at specific points in time with only the reward as feedback. This model is simulated to measure its performance, i.e. the increase in received reward after learning. Using this performance as baseline, we then simulate the model with various constraints imposed by the proposed implementation and compare the performance. The simulated constraints include discretized synaptic weights, a restricted interface between analog synapses and embedded processor, and mismatch of analog circuits. We find that probabilistic updates can increase the performance of low-resolution weights, a simple interface between analog synapses and processor is sufficient for learning, and performance is insensitive to mismatch. Further, we consider communication latency between wafer and the conventional control computer system that is simulating the environment. This latency increases the delay, with which the reward is sent to the embedded processor. Because of the time continuous operation of the analog synapses, delay can cause a deviation of the updates as compared to the not delayed situation. We find that for highly accelerated systems latency has to be kept to a minimum. This study demonstrates the suitability of the proposed implementation to emulate the selected reward modulated STDP learning rule.

q-bio.NC↗

arXiv2012

Nonnormal amplification in random balanced neuronal networks

In dynamical models of cortical networks, the recurrent connectivity can amplify the input given to the network in two distinct ways. One is induced by the presence of near-critical eigenvalues in the connectivity matrix W, producing large but slow activity fluctuations along the corresponding eigenvectors (dynamical slowing). The other relies on W being nonnormal, which allows the network activity to make large but fast excursions along specific directions. Here we investigate the tradeoff between nonnormal amplification and dynamical slowing in the spontaneous activity of large random neuronal networks composed of excitatory and inhibitory neurons. We use a Schur decomposition of W to separate the two amplification mechanisms. Assuming linear stochastic dynamics, we derive an exact expression for the expected amount of purely nonnormal amplification. We find that amplification is very limited if dynamical slowing must be kept weak. We conclude that, to achieve strong transient amplification with little slowing, the connectivity must be structured. We show that unidirectional connections between neurons of the same type together with reciprocal connections between neurons of different types, allow for amplification already in the fast dynamical regime. Finally, our results also shed light on the differences between balanced networks in which inhibition exactly cancels excitation, and those where inhibition dominates.

q-bio.NC↗

arXiv2010

Rescaling, thinning or complementing? On goodness-of-fit procedures for point process models and Generalized Linear Models

Generalized Linear Models (GLMs) are an increasingly popular framework for modeling neural spike trains. They have been linked to the theory of stochastic point processes and researchers have used this relation to assess goodness-of-fit using methods from point-process theory, e.g. the time-rescaling theorem. However, high neural firing rates or coarse discretization lead to a breakdown of the assumptions necessary for this connection. Here, we show how goodness-of-fit tests from point-process theory can still be applied to GLMs by constructing equivalent surrogate point processes out of time-series observations. Furthermore, two additional tests based on thinning and complementing point processes are introduced. They augment the instruments available for checking model adequacy of point processes as well as discretized models.

q-bio.NC↗

arXiv2005

Noise-enhanced computation in a model of a cortical column

Varied sensory systems use noise in order to enhance detection of weak signals. It has been conjectured in the literature that this effect, known as stochastic resonance, may take place in central cognitive processes such as the memory retrieval of arithmetical multiplication. We show in a simplified model of cortical tissue, that complex arithmetical calculations can be carried out and are enhanced in the presence of a stochastic background. The performance is shown to be positively correlated to the susceptibility of the network, defined as its sensitivity to a variation of the mean of its inputs. For nontrivial arithmetic tasks such as multiplication, stochastic resonance is an emergent property of the microcircuitry of the model network.

q-bio.NC↗

arXiv2005

Optimal Spike-Timing Dependent Plasticity for Precise Action Potential Firing

In timing-based neural codes, neurons have to emit action potentials at precise moments in time. We use a supervised learning paradigm to derive a synaptic update rule that optimizes via gradient ascent the likelihood of postsynaptic firing at one or several desired firing times. We find that the optimal strategy of up- and downregulating synaptic efficacies can be described by a two-phase learning window similar to that of Spike-Timing Dependent Plasticity (STDP). If the presynaptic spike arrives before the desired postsynaptic spike timing, our optimal learning rule predicts that the synapse should become potentiated. The dependence of the potentiation on spike timing directly reflects the time course of an excitatory postsynaptic potential. The presence and amplitude of depression of synaptic efficacies for reversed spike timing depends on how constraints are implemented in the optimization problem. Two different constraints, i.e., control of postsynaptic rates or control of temporal locality,are discussed.

q-bio.NC↗

arXiv2005

Transient Information Flow in a Network of Excitatory and Inhibitory Model Neurons: Role of Noise and Signal Autocorrelation

We investigate the performance of sparsely-connected networks of integrate-and-fire neurons for ultra-short term information processing. We exploit the fact that the population activity of networks with balanced excitation and inhibition can switch from an oscillatory firing regime to a state of asynchronous irregular firing or quiescence depending on the rate of external background spikes. We find that in terms of information buffering the network performs best for a moderate, non-zero, amount of noise. Analogous to the phenomenon of stochastic resonance the performance decreases for higher and lower noise levels. The optimal amount of noise corresponds to the transition zone between a quiescent state and a regime of stochastic dynamics. This provides a potential explanation on the role of non-oscillatory population activity in a simplified model of cortical micro-circuits.

q-bio.NC↗

arXiv2005

Signal buffering in random networks of spiking neurons: microscopic vs. macroscopic phenomena

In randomly connected networks of pulse-coupled elements a time-dependent input signal can be buffered over a short time. We studied the signal buffering properties in simulated networks as a function of the networks state, characterized by both the Lyapunov exponent of the microscopic dynamics and the macroscopic activity derived from mean-field theory. If all network elements receive the same signal, signal buffering over delays comparable to the intrinsic time constant of the network elements can be explained by macroscopic properties and works best at the phase transition to chaos. However, if only 20 percent of the network units receive a common time-dependent signal, signal buffering properties improve and can no longer be attributed to the macroscopic dynamics.

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