Slowness: An Objective for Spike-Timing-Dependent Plasticity?

Slow Feature Analysis (SFA) is an efficient algorithm for learning input-output functions that extract the most slowly varying features from a quickly varying signal. It has been successfully applied to the unsupervised learning of translation-, rotation-, and other invariances in a model of the visual system, to the learning of complex cell receptive fields, and, combined with a sparseness objective, to the self-organized formation of place cells in a model of the hippocampus. In order to arrive at a biologically more plausible implementation of this learning rule, we consider analytically how SFA could be realized in simple linear continuous and spiking model neurons. It turns out that for the continuous model neuron SFA can be implemented by means of a modified version of standard Hebbian learning. In this framework we provide a connection to the trace learning rule for invariance learning. We then show that for Poisson neurons spike-timing-dependent plasticity (STDP) with a specific learning window can learn the same weight distribution as SFA. Surprisingly, we find that the appropriate learning rule reproduces the typical STDP learning window. The shape as well as the timescale are in good agreement with what has been measured experimentally. This offers a completely novel interpretation for the functional role of spike-timing-dependent plasticity in physiological neurons.

Biologically plausible unsupervised learning in shallow and deep neural networks

Bernd Albert Illing

The way our brain learns to disentangle complex signals into unambiguous concepts is fascinating but remains largely unknown. There is evidence, however, that hierarchical neural representations play a key role in the cortex. This thesis investigates biologically plausible models of unsupervised learning of hierarchical representations as found in the brain and modern computer vision models. We use computational modeling to address three main questions at the intersection of artificial intelligence (AI) and computational neuroscience.The first question is: What are useful neural representations and when are deep hierarchical representations needed? We approach this point with a systematic study of biologically plausible unsupervised feature learning in a shallow 2-layer networks on digit (MNIST) and object (CIFAR10) classification. Surprisingly, random features support high performance, especially for large hidden layers. When combined with localized receptive fields, random feature networks approach the performance of supervised backpropagation on MNIST, but not on CIFAR10. We suggest that future models of biologically plausible learning should outperform such random feature benchmarks on MNIST, or that such models should be evaluated in different ways.The second question is: How can hierarchical representations be learned with mechanisms supported by neuroscientific evidence? We cover this question by proposing a unifying Hebbian model, inspired by common models of V1 simple and complex cells based on unsupervised sparse coding and temporal invariance learning. In shallow 2-layer networks, our model reproduces learning of simple and complex cell receptive fields, as found in V1. In deeper networks, we stack multiple layers of Hebbian learning but find that it does not yield hierarchical representations of increasing usefulness. From this, we hypothesise that standard Hebbian rules are too constrained to build increasingly useful representations, as observed in higher areas of the visual cortex or deep artificial neural networks.The third question is: Can AI inspire learning models that build deep representations and are still biologically plausible? We address this question by proposing a learning rule that takes inspiration from neuroscience and recent advances in self-supervised deep learning. The proposed rule is Hebbian, i.e. only depends on pre- and post-synaptic neuronal activity, but includes additional local factors, namely predictive dendritic input and widely broadcasted modulation factors. Algorithmically, this rule applies self-supervised contrastive predictive learning to a causal, biological setting using saccades. We find that networks trained with this generalised Hebbian rule build deep hierarchical representations of images, speech and video.We see our modeling as a potential starting point for both, new hypotheses, that can be tested experimentally, and novel AI models that could benefit from added biological realism.

EPFL2021

Theory of representation learning in cortical neural networks

Carlos Stein Naves de Brito

Our brain continuously self-organizes to construct and maintain an internal representation of the world based on the information arriving through sensory stimuli. Remarkably, cortical areas related to different sensory modalities appear to share the same functional unit, the neuron, and develop through the same learning mechanism, synaptic plasticity. It motivates the conjecture of a unifying theory to explain cortical representational learning across sensory modalities. In this thesis we present theories and computational models of learning and optimization in neural networks, postulating functional properties of synaptic plasticity that support the apparent universal learning capacity of cortical networks. In the past decades, a variety of theories and models have been proposed to describe receptive field formation in sensory areas. They include normative models such as sparse coding, and bottom-up models such as spike-timing dependent plasticity. We bring together candidate explanations by demonstrating that in fact a single principle is sufficient to explain receptive field development. First, we show that many representative models of sensory development are in fact implementing variations of a common principle: nonlinear Hebbian learning. Second, we reveal that nonlinear Hebbian learning is sufficient for receptive field formation through sensory inputs. A surprising result is that our findings are independent of specific details, and allow for robust predictions of the learned receptive fields. Thus nonlinear Hebbian learning and natural statistics can account for many aspects of receptive field formation across models and sensory modalities. The Hebbian learning theory substantiates that synaptic plasticity can be interpreted as an optimization procedure, implementing stochastic gradient descent. In stochastic gradient descent inputs arrive sequentially, as in sensory streams. However, individual data samples have very little information about the correct learning signal, and it becomes a fundamental problem to know how many samples are required for reliable synaptic changes. Through estimation theory, we develop a novel adaptive learning rate model, that adapts the magnitude of synaptic changes based on the statistics of the learning signal, enabling an optimal use of data samples. Our model has a simple implementation and demonstrates improved learning speed, making this a promising candidate for large artificial neural network applications. The model also makes predictions on how cortical plasticity may modulate synaptic plasticity for optimal learning. The optimal sampling size for reliable learning allows us to estimate optimal learning times for a given model. We apply this theory to derive analytical bounds on times for the optimization of synaptic connections. First, we show this optimization problem to have exponentially many saddle-nodes, which lead to small gradients and slow learning. Second, we show that the number of input synapses to a neuron modulates the magnitude of the initial gradient, determining the duration of learning. Our final result reveals that the learning duration increases supra-linearly with the number of synapses, suggesting an effective limit on synaptic connections and receptive field sizes in developing neural networks.

EPFL2016

Biologically plausible unsupervised learning in shallow and deep neural networks

Bernd Albert Illing

EPFL2021

Theory of representation learning in cortical neural networks

Carlos Stein Naves de Brito

EPFL2016