Neural correlates of learning and memory formation have been reported at different stages of the olfactory pathway in both vertebrates and invertebrates. However, the contribution of different neurons to the formation of a memory trace is little understood. Mushroom bodies (MBs) in the insect brain are higher-order structures involved in integration of olfactory, visual, and mechanosensory information and in memory formation. Here we focus on the ensemble spiking activity of single MB output neurons (ENs) when honeybees learned to associate an odor with reward. A large group of ENs (ϳ50%) changed their odor response spectra by losing or gaining sensitivity for specific odors. This response switching was dominated by the rewarded stimulus (CSϩ), which evoked exclusively recruitment. The remaining ENs did not change their qualitative odor spectrum but modulated their tuning strength, again dominated by increased responses to the CSϩ. While the bees showed a conditioned response (proboscis extension) after a few acquisition trials, no short-term effects were observed in the neuronal activity. In both EN types, associative plastic changes occurred only during retention 3 h after conditioning. Thus, long-term but not short-term memory was reflected by increased EN activity to the CSϩ. During retention, the EN ensemble separated the CSϩ most differently from the CSϪ and control odors ϳ140 ms after stimulus onset. The learned behavioral response appeared ϳ330 ms later. It is concluded that after memory consolidation, the ensemble activity of the MB output neurons predicts the meaning of the stimulus (reward) and may provide the prerequisite for the expression of the learned behavior.
Computational neuroscience has uncovered a number of computational principles used by nervous systems. At the same time, neuromorphic hardware has matured to a state where fast silicon implementations of complex neural networks have become feasible. En route to future technical applications of neuromorphic computing the current challenge lies in the identification and implementation of functional brain algorithms. Taking inspiration from the olfactory system of insects, we constructed a spiking neural network for the classification of multivariate data, a common problem in signal and data analysis. In this model, real-valued multivariate data are converted into spike trains using "virtual receptors" (VRs). Their output is processed by lateral inhibition and drives a winner-take-all circuit that supports supervised learning. VRs are conveniently implemented in software, whereas the lateral inhibition and classification stages run on accelerated neuromorphic hardware. When trained and tested on real-world datasets, we find that the classification performance is on par with a naïve Bayes classifier. An analysis of the network dynamics shows that stable decisions in output neuron populations are reached within less than 100 ms of biological time, matching the time-to-decision reported for the insect nervous system. Through leveraging a population code, the network tolerates the variability of neuronal transfer functions and trial-totrial variation that is inevitably present on the hardware system. Our work provides a proof of principle for the successful implementation of a functional spiking neural network on a configurable neuromorphic hardware system that can readily be applied to realworld computing problems.bioinspired computing | spiking networks | machine learning | multivariate classification T he remarkable sensory and behavioral capabilities of all higher organisms are provided by the network of neurons in their nervous systems. The computing principles of the brain have inspired many powerful algorithms for data processing, most importantly the perceptron and, building on top of that, multilayer artificial neural networks, which are being applied with great success to various data analysis problems (1). Although these networks operate with continuous values, computation in biological neuronal networks relies on the exchange of action potentials, or "spikes."Simulating networks of spiking neurons with software tools is computationally intensive, imposing limits to the duration of simulations and maximum network size. To overcome this limitation, several groups around the world have started to develop hardware realizations of spiking neuron models and neuronal networks (2-10) for studying the behavior of biological networks (11). The approach of the Spikey hardware system used in the present study is to enable high-throughput network simulations by speeding up computation by a factor of 10 4 compared with biological real time (12, 13). It has been developed as a reconfigurable multineuron computing substrat...
We present a method to estimate the neuronal firing rate from single-trial spike trains. The method, based on convolution of the spike train with a fixed kernel function, is calibrated by means of simulated spike trains for a representative selection of realistic dynamic rate functions. We derive rules for the optimized use and performance of the kernel method, specifically with respect to an effective choice of the shape and width of the kernel functions. An application of our technique to the on-line, single-trial reconstruction of arm movement trajectories from multiple single-unit spike trains using dynamic population vectors illustrates a possible use of the proposed method.
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