Despite being originally inspired by the central nervous system, artificial neural networks have diverged from their biological archetypes as they have been remodeled to fit particular tasks. In this paper, we review several possibilites to reverse map these architectures to biologically more realistic spiking networks with the aim of emulating them on fast, low-power neuromorphic hardware. Since many of these devices employ analog components, which cannot be perfectly controlled, finding ways to compensate for the resulting effects represents a key challenge. Here, we discuss three different strategies to address this problem: the addition of auxiliary network components for stabilizing activity, the utilization of inherently robust architectures and a training method for hardware-emulated networks that functions without perfect knowledge of the system's dynamics and parameters. For all three scenarios, we corroborate our theoretical considerations with experimental results on accelerated analog neuromorphic platforms.

We reexamine in this article the conceptual and mathematical framework for understanding the organization of plasticity in spiking neural networks. We propose that inherent stochasticity enables synaptic plasticity to carry out probabilistic inference by sampling from a posterior distribution of synaptic parameters. This view provides a viable alternative to existing models that propose convergence of synaptic weights to maximum likelihood parameters. It explains how priors on weight distributions and connection probabilities can be merged optimally with learned experience. In simulations we show that our model for synaptic plasticity allows spiking neural networks to compensate continuously for unforeseen disturbances. Furthermore it provides a normative mathematical framework to better understand the permanent variability and rewiring observed in brain networks.

The control of neuroprosthetic devices from the activity of motor cortex neurons benefits from learning effects where the function of these neurons is adapted to the control task. It was recently shown that tuning properties of neurons in monkey motor cortex are adapted selectively in order to compensate for an erroneous interpretation of their activity. In particular, it was shown that the tuning curves of those neurons whose preferred directions had been misinterpreted changed more than those of other neurons. In this article, we show that the experimentally observed self-tuning properties of the system can be explained on the basis of a simple learning rule. This learning rule utilizes neuronal noise for exploration and performs Hebbian weight updates that are modulated by a global reward signal. In contrast to most previously proposed reward-modulated Hebbian learning rules, this rule does not require extraneous knowledge about what is noise and what is signal. The learning rule is able to optimize the performance of the model system within biologically realistic periods of time and under high noise levels. When the neuronal noise is fitted to experimental data, the model produces learning effects similar to those found in monkey experiments.

Many models for computations in recurrent networks of neurons assume that the network state moves from some initial state to some fixed point attractor or limit cycle that represents the output of the computation. However experimental data show that in response to a sensory stimulus the network state moves from its initial state through a trajectory of network states and eventually returns to the initial state, without reaching an attractor or limit cycle in between. This type of network response, where salient information about external stimuli is encoded in characteristic trajectories of continuously varying network states, raises the question how a neural system could compute with such code, and arrive for example at a temporally stable classification of the external stimulus. We show that a known unsupervised learning algorithm, Slow Feature Analysis (SFA), could be an important ingredient for extracting stable information from these network trajectories. In fact, if sensory stimuli are more often followed by another stimulus from the same class than by a stimulus from another class, SFA approaches the classification capability of Fishers Linear Discriminant (FLD), a powerful algorithm for supervised learning. We apply this principle to simulated cortical microcircuits, and show that it enables readout neurons to learn discrimination of spoken digits and detection of repeating firing patterns within a stream of spike trains with the same firing statistics, without requiring any supervision for learning.

The principles by which spiking neurons contribute to the astounding computational power of generic cortical microcircuits, and how spike-timing-dependent plasticity (STDP) of synaptic weights could generate and maintain this computational function, are unknown. We show here that STDP, in conjunction with a stochastic soft winner-take-all (WTA) circuit, induces spiking neurons to generate through their synaptic weights implicit internal models for subclasses (or causes") of the high-dimensional spike patterns of hundreds of pre-synaptic neurons. Hence these neurons will fire after learning whenever the current input best matches their internal model. The resulting computational function of soft WTA circuits, a common network motif of cortical microcircuits, could therefore be a drastic dimensionality reduction of information streams, together with the autonomous creation of internal models for the probability distributions of their input patterns. We show that the autonomous generation and maintenance of this computational function can be explained on the basis of rigorous mathematical principles. In particular, we show that STDP is able to approximate a stochastic online Expectation-Maximization (EM) algorithm for modeling the input data. A corresponding result is shown for Hebbian learning in artificial neural networks."

Uncertainty is omnipresent when we perceive or interact with our environment, and the Bayesian framework provides computational methods for dealing with it. Mathematical models for Bayesian decision making typically require datastructures that are hard to implement in neural networks. This article shows that even the simplest and experimentally best supported type of synaptic plasticity, Hebbian learning, in combination with a sparse, redundant neural code, can in principle learn to infer optimal Bayesian decisions. We present a concrete Hebbian learning rule operating on log-probability ratios. Modulated by reward-signals, this Hebbian plasticity rule also provides a new perspective for understanding how Bayesian inference could support fast reinforcement learning in the brain. In particular we show that recent experimental results by Yang and Shadlen [1] on reinforcement learning of probabilistic inference in primates can be modeled in this way.

Reward-modulated spike-timing-dependent plasticity (STDP) has recently emerged as a candidate for a learning rule that could explain how local learning rules at single synapses support behaviorally relevant adaptive changes in complex networksof spiking neurons. However the potential and limitations of this learning rule could so far only be tested through computer simulations. This article providestools for an analytic treatment of reward-modulated STDP, which allow us to predict under which conditions reward-modulated STDP will be able to achieve a desired learning effect. In particular, we can produce in this way a theoretical explanation and a computer model for a fundamental experimental finding on biofeedback in monkeys (reported in [1]).

We show that under suitable assumptions (primarily linearization) a simple and perspicuous online learning rule for Information Bottleneck optimization with spiking neurons can be derived. This rule performs on common benchmark tasks as well as a rather complex rule that has previously been proposed \cite{KlampflETAL:07b}. Furthermore, the transparency of this new learning rule makes a theoretical analysis of its convergence properties feasible. A variation of this learning rule (with sign changes) provides a theoretically founded method for performing Principal Component Analysis {(PCA)} with spiking neurons. By applying this rule to an ensemble of neurons, different principal components of the input can be extracted. In addition, it is possible to preferentially extract those principal components from incoming signals $X$ that are related or are not related to some additional target signal $Y_T$. In a biological interpretation, this target signal $Y_T$ (also called relevance variable) could represent proprioceptive feedback, input from other sensory modalities, or top-down signals.

We use multi-electrode recordings from cat primary visual cortex and investigate whether a simple linear classifier can extract information about the presented stimuli. We find that information is extractable and that it even lasts for several hundred milliseconds after the stimulus has been removed. In a fast sequence of stimulus presentation, information about both new and old stimuli is present simultaneously and nonlinear relations between these stimuli can be extracted. These results suggest nonlinear properties of cortical representations. The important implications of these properties for the nonlinear brain theory are discussed.

The extraction of statistically independent components from high-dimensional multi-sensory input streams is assumed to be an essential component of sensory processing in the brain. Such independent component analysis (or blind source separation) could provide a less redundant representation of information about the external world. Another powerful processing strategy is to extract preferentially those components from high-dimensional input streams that are related to other information sources, such as internal predictions or proprioceptive feedback. This strategy allows the optimization of internal representation according to the information bottleneckmethod. However, concrete learning rules that implement these general unsupervised learning principles for spiking neurons are still missing. We show how both information bottleneck optimization and the extraction of independent componentscan in principle be implemented with stochastically spiking neurons with refractoriness. The new learning rule that achieves this is derived from abstract information optimization principles.