SuperSpike: Supervised learning in multi-layer spiking neural networks

A vast majority of computation in the brain is performed by spiking neural networks. Despite the ubiquity of such spiking, we currently lack an understanding of how biological spiking neural circuits learn and compute in-vivo, as well as how we can instantiate such capabilities in artificial spiking circuits in-silico. Here we revisit the problem of supervised learning in temporally coding multi-layer spiking neural networks. First, by using a surrogate gradient approach, we derive SuperSpike, a nonlinear voltage-based three factor learning rule capable of training multi-layer networks of deterministic integrate-and-fire neurons to perform nonlinear computations on spatiotemporal spike patterns. Second, inspired by recent results on feedback alignment, we compare the performance of our learning rule under different credit assignment strategies for propagating output errors to hidden units. Specifically, we test uniform, symmetric and random feedback, finding that simpler tasks can be solved with any type of feedback, while more complex tasks require symmetric feedback. In summary, our results open the door to obtaining a better scientific understanding of learning and computation in spiking neural networks by advancing our ability to train them to solve nonlinear problems involving transformations between different spatiotemporal spike-time patterns

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

Logarithmic distributions prove that intrinsic learning is Hebbian

In this paper, we present data for the lognormal distributions of spike rates, synaptic weights and intrinsic excitability (gain) for neurons in various brain areas, such as auditory or visual cortex, hippocampus, cerebellum, striatum, midbrain nuclei. We find a remarkable consistency of heavy-tailed, specifically lognormal, distributions for rates, weights and gains in all brain areas examined. The difference between strongly recurrent and feed-forward connectivity (cortex vs. striatum and cerebellum), neurotransmitter (GABA (striatum) or glutamate (cortex)) or the level of activation (low in cortex, high in Purkinje cells and midbrain nuclei) turns out to be irrelevant for this feature. Logarithmic scale distribution of weights and gains appears to be a general, functional property in all cases analyzed. We then created a generic neural model to investigate adaptive learning rules that create and maintain lognormal distributions. We conclusively demonstrate that not only weights, but also intrinsic gains, need to have strong Hebbian learning in order to produce and maintain the experimentally attested distributions. This provides a solution to the long-standing question about the type of plasticity exhibited by intrinsic excitability

Role of homeostasis in learning sparse representations

Neurons in the input layer of primary visual cortex in primates develop edge-like receptive fields. One approach to understanding the emergence of this response is to state that neural activity has to efficiently represent sensory data with respect to the statistics of natural scenes. Furthermore, it is believed that such an efficient coding is achieved using a competition across neurons so as to generate a sparse representation, that is, where a relatively small number of neurons are simultaneously active. Indeed, different models of sparse coding, coupled with Hebbian learning and homeostasis, have been proposed that successfully match the observed emergent response. However, the specific role of homeostasis in learning such sparse representations is still largely unknown. By quantitatively assessing the efficiency of the neural representation during learning, we derive a cooperative homeostasis mechanism that optimally tunes the competition between neurons within the sparse coding algorithm. We apply this homeostasis while learning small patches taken from natural images and compare its efficiency with state-of-the-art algorithms. Results show that while different sparse coding algorithms give similar coding results, the homeostasis provides an optimal balance for the representation of natural images within the population of neurons. Competition in sparse coding is optimized when it is fair. By contributing to optimizing statistical competition across neurons, homeostasis is crucial in providing a more efficient solution to the emergence of independent components

Storage of phase-coded patterns via STDP in fully-connected and sparse network: a study of the network capacity

We study the storage and retrieval of phase-coded patterns as stable dynamical attractors in recurrent neural networks, for both an analog and a integrate-and-fire spiking model. The synaptic strength is determined by a learning rule based on spike-time-dependent plasticity, with an asymmetric time window depending on the relative timing between pre- and post-synaptic activity. We store multiple patterns and study the network capacity. For the analog model, we find that the network capacity scales linearly with the network size, and that both capacity and the oscillation frequency of the retrieval state depend on the asymmetry of the learning time window. In addition to fully-connected networks, we study sparse networks, where each neuron is connected only to a small number z << N of other neurons. Connections can be short range, between neighboring neurons placed on a regular lattice, or long range, between randomly chosen pairs of neurons. We find that a small fraction of long range connections is able to amplify the capacity of the network. This imply that a small-world-network topology is optimal, as a compromise between the cost of long range connections and the capacity increase. Also in the spiking integrate and fire model the crucial result of storing and retrieval of multiple phase-coded patterns is observed. The capacity of the fully-connected spiking network is investigated, together with the relation between oscillation frequency of retrieval state and window asymmetry