Biologically plausible deep learning -- but how far can we go with shallow networks?

Training deep neural networks with the error backpropagation algorithm is considered implausible from a biological perspective. Numerous recent publications suggest elaborate models for biologically plausible variants of deep learning, typically defining success as reaching around 98% test accuracy on the MNIST data set. Here, we investigate how far we can go on digit (MNIST) and object (CIFAR10) classification with biologically plausible, local learning rules in a network with one hidden layer and a single readout layer. The hidden layer weights are either fixed (random or random Gabor filters) or trained with unsupervised methods (PCA, ICA or Sparse Coding) that can be implemented by local learning rules. The readout layer is trained with a supervised, local learning rule. We first implement these models with rate neurons. This comparison reveals, first, that unsupervised learning does not lead to better performance than fixed random projections or Gabor filters for large hidden layers. Second, networks with localized receptive fields perform significantly better than networks with all-to-all connectivity and can reach backpropagation performance on MNIST. We then implement two of the networks - fixed, localized, random & random Gabor filters in the hidden layer - with spiking leaky integrate-and-fire neurons and spike timing dependent plasticity to train the readout layer. These spiking models achieve > 98.2% test accuracy on MNIST, which is close to the performance of rate networks with one hidden layer trained with backpropagation. The performance of our shallow network models is comparable to most current biologically plausible models of deep learning. Furthermore, our results with a shallow spiking network provide an important reference and suggest the use of datasets other than MNIST for testing the performance of future models of biologically plausible deep learning.Comment: 14 pages, 4 figure

Neural principles underlying motor learning and adaptation

Animals, and especially humans, can learn to flexibly adjust their movements to changing environments. The neural principles underlying this remarkable capability are still not fully understood. Among the most prominent brain regions controlling movement is primary motor cortex (M1). Adapted motor behaviour can be related to a change in neural activity within this region. Yet, the rules guiding this activity change, and thus behavioural adaptation, remain unclear. The overall aim of this thesis is to investigate the learning process(es) governing the described change in activity in M1 and, with that, the change in behaviour. Computational modelling is used to study three specific aspects of learning: 1. What constrains learning to favour some neural activity patterns over others? 2. Can we identify where in a hierarchical pathway learning is happening? 3. How can sensory feedback guide the learning process? We start by investigating what kind of biological constraints differentially affect learning of new neural activity that either preserves coactivation patterns between neurons (within-manifold learning), or requires learning of new coactivation patterns (outside-manifold learning). We propose a new explanation - the learnability of feedback signals - for why within-manifold activity patterns can be easier learned than outside-manifold activity patterns. In the second part we develop a hierarchical model of the motor system to investigate whether we can derive where learning has happened from only measuring neural activity. Lastly, we investigate how the brain could implement a biologically plausible learning rule which allows it to correctly assign errors and update recurrent connectivity in a goal-driven manner. Overall, our work offers new perspectives on the role of M1 for motor learning and adaptation, challenges current beliefs, and puts a focus on the role of feedback signals for local plasticity in M1.Open Acces

Computational roles of cortico-cerebellar loops in temporal credit assignment

Animal survival depends on behavioural adaptation to the environment. This is thought to be enabled by plasticity in the neural circuit. However, the laws which govern neural plasticity are unclear. From a functional aspect, it is desirable to correctly identify, or assign “credit” for, the neurons or synapses responsible for the task decision and subsequent performance. In the biological circuit, the intricate, non-linear interactions involved in neural networks makes appropriately assigning credit to neurons highly challenging. In the temporal domain, this is known as the temporal credit assignment (TCA) problem. This Thesis considers the role the cerebellum – a powerful subcortical structure with strong error-guided plasticity rules – as a solution to TCA in the brain. In particular, I use artificial neural networks as a means to model and understand the mechanisms by which the cerebellum can support learning in the neocortex via the cortico-cerebellar loop. I introduce two distinct but compatible computational models of cortico-cerebellar interaction. The first model asserts that the cerebellum provides the neocortex predictive feedback, modeled in the form of error gradients, with respect to its current activity. This predictive feedback enables better credit assignment in the neocortex and effectively removes the lock between feedforward and feedback processing in cortical networks. This model captures observed long-term deficits associated with cerebellar dysfunction, namely cerebellar dysmetria, in both the motor and non-motor domain. Predictions are also made with respect to alignment of cortico-cerebellar activity during learning and the optimal task conditions for cerebellar contribution. The second model also looks at the role of the cerebellum in learning, but now considers its ability to instantaneously drive the cortex towards desired task dynamics. Unlike the first model, this model does not assume any local cortical plasticity need take place at all and task-directed learning can effectively be outsourced to the cerebellum. This model captures recent optogenetic studies in mice which show the cerebellum as a necessary component for the maintenance of desired cortical dynamics and ensuing behaviour. I also show that this driving input can eventually be used as a teaching signal for the cortical circuit, thereby conceptually unifying the two models. Overall, this Thesis explores the computational role of the cerebellum and cortico-cerebellar loops for task acquisition and maintenance in the brain

Visual motion processing and human tracking behavior

The accurate visual tracking of a moving object is a human fundamental skill that allows to reduce the relative slip and instability of the object's image on the retina, thus granting a stable, high-quality vision. In order to optimize tracking performance across time, a quick estimate of the object's global motion properties needs to be fed to the oculomotor system and dynamically updated. Concurrently, performance can be greatly improved in terms of latency and accuracy by taking into account predictive cues, especially under variable conditions of visibility and in presence of ambiguous retinal information. Here, we review several recent studies focusing on the integration of retinal and extra-retinal information for the control of human smooth pursuit.By dynamically probing the tracking performance with well established paradigms in the visual perception and oculomotor literature we provide the basis to test theoretical hypotheses within the framework of dynamic probabilistic inference. We will in particular present the applications of these results in light of state-of-the-art computer vision algorithms