arXiv subjects

Walter Senn

Publications and source records attributed to Walter Senn.

At least 19 recordsLinked to original sources

arXiv2026

Dendritic structure enables powerful plasticity

Over the past decades, it has become increasingly clear that the complex morphology of cortical neurons is more than just a quirk of evolution, and that dendritic compartments serve as computational elements in their own right, rather than just providing connections between nerve cell bodies. While most computational studies discuss the enhanced representational capabilities of multi-compartment models as compared to point neurons, we focus here on the implications of neuronal morphology for synaptic plasticity. We argue that the ability of single neurons to simultaneously encode multiple pieces of information gives synapses local access to more than just the classical Hebbian pre- and postsynaptic terms, and with much greater specificity and reaction speed than permitted by other globally modulated factors. Based on a comparative review of recent dendritic learning models, we show how such neuronal compartmentalization can provide synapses with the means for calculating various forms of error signals, which in turn give rise to powerful real-time and fully local instantiations of deep learning through gradient descent. Implemented within cortical microcircuits capable of propagating and manipulating these errors, compartmentalized neurons thus ultimately enable the learning of far more complex tasks than are achievable by globally modulated Hebbian plasticity alone.

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arXiv2026

A Variational Latent Equilibrium for Learning in Neuronal Circuits

Brains remain unrivaled in their ability to recognize and generate complex spatiotemporal patterns. While AI is able to reproduce some of these capabilities, deep learning algorithms remain largely at odds with our current understanding of brain circuitry and dynamics. This is prominently the case for backpropagation through time (BPTT), the go-to algorithm for learning complex temporal dependencies. In this work we propose a general formalism to approximate BPTT in a controlled, biologically plausible manner. Our approach builds on, unifies and extends several previous approaches to local, time-continuous, phase-free spatiotemporal credit assignment based on principles of energy conservation and extremal action. Our starting point is a prospective energy function of neuronal states, from which we calculate real-time error dynamics for time-continuous neuronal networks. In the general case, this provides a simple and straightforward derivation of the adjoint method result for neuronal networks, the time-continuous equivalent to BPTT. With a few modifications, we can turn this into a fully local (in space and time) set of equations for neuron and synapse dynamics. Our theory provides a rigorous framework for spatiotemporal deep learning in the brain, while simultaneously suggesting a blueprint for physical circuits capable of carrying out these computations. These results reframe and extend the recently proposed Generalized Latent Equilibrium (GLE) model.

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arXiv2025

Teaching signal synchronization in deep neural networks with prospective neurons

Working memory requires the brain to maintain information from the recent past to guide ongoing behavior. Neurons can contribute to this capacity by slowly integrating their inputs over time, creating persistent activity that outlasts the original stimulus. However, when these slowly integrating neurons are organized hierarchically, they introduce cumulative delays that create a fundamental challenge for learning: teaching signals that indicate whether behavior was correct or incorrect arrive out-of-sync with the neural activity they are meant to instruct. Here, we demonstrate that neurons enhanced with an adaptive current can compensate for these delays by responding to external stimuli prospectively -- effectively predicting future inputs to synchronize with them. First, we show that such prospective neurons enable teaching signal synchronization across a range of learning algorithms that propagate error signals through hierarchical networks. Second, we demonstrate that this successfully guides learning in slowly integrating neurons, enabling the formation and retrieval of memories over extended timescales. We support our findings with a mathematical analysis of the prospective coding mechanism and learning experiments on motor control tasks. Together, our results reveal how neural adaptation could solve a critical timing problem and enable efficient learning in dynamic environments.

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arXiv2025

Neural mechanisms of predictive processing: a collaborative community experiment through the OpenScope program

This review synthesizes advances in predictive processing within the sensory cortex. Predictive processing theorizes that the brain continuously predicts sensory inputs, refining neuronal responses by highlighting prediction errors. We identify key computational primitives, such as stimulus adaptation, dendritic computation, excitatory/inhibitory balance and hierarchical processing, as central to this framework. Our review highlights convergences, such as top-down inputs and inhibitory interneurons shaping mismatch signals, and divergences, including species-specific hierarchies and modality-dependent layer roles. To address these conflicts, we propose experiments in mice and primates using in-vivo two-photon imaging and electrophysiological recordings to test whether temporal, motor, and omission mismatch stimuli engage shared or distinct mechanisms. The resulting dataset, collected and shared via the OpenScope program, will enable model validation and community analysis, fostering iterative refinement and refutability to decode the neural circuits of predictive processing.

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arXiv2025

Multihead self-attention in cortico-thalamic circuits

Both biological cortico-thalamic networks and artificial transformer networks use canonical computations to perform a wide range of cognitive tasks. In this work, we propose that the structure of cortico-thalamic circuits is well suited to realize a computation analogous to multihead self-attention, the main algorithmic innovation of transformer networks. We assign distinct computational roles to superficial and deep pyramidal cells of the cortex: while superficial pyramidal cells maintain a key-value memory, deep pyramidal cells encode the current query, gain-modulated by the key-value memory in the superficial layer. We show that the structure of this computation matches the fine-grained structure of core and matrix projections from the thalamus to the cortex. We then suggest the parallel between one head of attention and a cortical area, and propose that a thalamo-cortico-thalamic pathway implements a computation akin to a multihead, unnormalized, linear self-attention block. Cross-attention corresponds to the key-value memory of one cortical area being used for retrieval by the query in another cortical area. Finally, as a first step towards a mechanistic theory of synaptic learning of cortical transformers, we derive the formal gradients of a typical loss function with respect to the parameters of such computation.

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arXiv2025

Building functional and mechanistic models of cortical computation based on canonical cell type connectivity

Neuronal circuits of the cerebral cortex are the structural basis of mammalian cognition. The same qualitative components and connectivity motifs are repeated across functionally specialized cortical areas and mammalian species, suggesting a single underlying algorithmic motif. Here, we propose a perspective on current knowledge of the cortical structure, from which we extract two core principles for computational modeling. The first principle is that cortical cell types fulfill distinct computational roles. The second principle is that cortical connectivity can be efficiently characterized by only a few canonical blueprints of connectivity between cell types. Starting with these two foundational principles, we outline a general framework for building functional and mechanistic models of cortical circuits.

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arXiv2024

Prospective and retrospective coding in cortical neurons

Brains can process sensory information from different modalities at astonishing speed; this is surprising as the integration of inputs through the membrane of each individual neuron already causes a delayed response. Neuronal recordings {\em in vitro} reveal a possible explanation for this fast processing, in terms of individual neurons advancing their output firing rates with respect to the input, a concept which we refer to as prospective coding. The underlying mechanisms of prospective coding, however, are not completely understood. We propose a mechanistic explanation for individual neurons advancing their output on the level of single action potentials and instantaneous firing rates. We show that the spike generation mechanism can be the source for prospective (advanced) or retrospective (delayed) responses. A simplified Hodgkin-Huxley model identifies sodium inactivation as a source for prospective firing, controlling the timing of the neuron's output as a function of the voltage and its temporal derivative. We further show that slow adaptation processes, such as spike-frequency adaptation or deactivating dendritic currents, represent mechanisms generating prospective firing for inputs that undergo slow temporal modulations. In general, we show that adaptation processes at different time scales can cause advanced neuronal responses to time-varying inputs that are modulated on the corresponding time scales.

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arXiv2024

ELiSe: Efficient Learning of Sequences in Structured Recurrent Networks

Behavior can be described as a temporal sequence of actions driven by neural activity. To learn complex sequential patterns in neural networks, memories of past activities need to persist on significantly longer timescales than the relaxation times of single-neuron activity. While recurrent networks can produce such long transients, training these networks is a challenge. Learning via error propagation confers models such as FORCE, RTRL or BPTT a significant functional advantage, but at the expense of biological plausibility. While reservoir computing circumvents this issue by learning only the readout weights, it does not scale well with problem complexity. We propose that two prominent structural features of cortical networks can alleviate these issues: the presence of a certain network scaffold at the onset of learning and the existence of dendritic compartments for enhancing neuronal information storage and computation. Our resulting model for Efficient Learning of Sequences (ELiSe) builds on these features to acquire and replay complex non-Markovian spatio-temporal patterns using only local, always-on and phase-free synaptic plasticity. We showcase the capabilities of ELiSe in a mock-up of birdsong learning, and demonstrate its flexibility with respect to parametrization, as well as its robustness to external disturbances.

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arXiv2024

Learning efficient backprojections across cortical hierarchies in real time

Models of sensory processing and learning in the cortex need to efficiently assign credit to synapses in all areas. In deep learning, a known solution is error backpropagation, which however requires biologically implausible weight transport from feed-forward to feedback paths. We introduce Phaseless Alignment Learning (PAL), a bio-plausible method to learn efficient feedback weights in layered cortical hierarchies. This is achieved by exploiting the noise naturally found in biophysical systems as an additional carrier of information. In our dynamical system, all weights are learned simultaneously with always-on plasticity and using only information locally available to the synapses. Our method is completely phase-free (no forward and backward passes or phased learning) and allows for efficient error propagation across multi-layer cortical hierarchies, while maintaining biologically plausible signal transport and learning. Our method is applicable to a wide class of models and improves on previously known biologically plausible ways of credit assignment: compared to random synaptic feedback, it can solve complex tasks with less neurons and learn more useful latent representations. We demonstrate this on various classification tasks using a cortical microcircuit model with prospective coding.

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arXiv2023

Learning beyond sensations: how dreams organize neuronal representations

Semantic representations in higher sensory cortices form the basis for robust, yet flexible behavior. These representations are acquired over the course of development in an unsupervised fashion and continuously maintained over an organism's lifespan. Predictive learning theories propose that these representations emerge from predicting or reconstructing sensory inputs. However, brains are known to generate virtual experiences, such as during imagination and dreaming, that go beyond previously experienced inputs. Here, we suggest that virtual experiences may be just as relevant as actual sensory inputs in shaping cortical representations. In particular, we discuss two complementary learning principles that organize representations through the generation of virtual experiences. First, "adversarial dreaming" proposes that creative dreams support a cortical implementation of adversarial learning in which feedback and feedforward pathways engage in a productive game of trying to fool each other. Second, "contrastive dreaming" proposes that the invariance of neuronal representations to irrelevant factors of variation is acquired by trying to map similar virtual experiences together via a contrastive learning process. These principles are compatible with known cortical structure and dynamics and the phenomenology of sleep thus providing promising directions to explain cortical learning beyond the classical predictive learning paradigm.

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arXiv2023

Confidence and second-order errors in cortical circuits

Minimization of cortical prediction errors has been considered a key computational goal of the cerebral cortex underlying perception, action and learning. However, it is still unclear how the cortex should form and use information about uncertainty in this process. Here, we formally derive neural dynamics that minimize prediction errors under the assumption that cortical areas must not only predict the activity in other areas and sensory streams but also jointly project their confidence (inverse expected uncertainty) in their predictions. In the resulting neuronal dynamics, the integration of bottom-up and top-down cortical streams is dynamically modulated based on confidence in accordance with the Bayesian principle. Moreover, the theory predicts the existence of cortical second-order errors, comparing confidence and actual performance. These errors are propagated through the cortical hierarchy alongside classical prediction errors and are used to learn the weights of synapses responsible for formulating confidence. We propose a detailed mapping of the theory to cortical circuitry, discuss entailed functional interpretations and provide potential directions for experimental work.

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arXiv2023

Conductance-based dendrites perform Bayes-optimal cue integration

A fundamental function of cortical circuits is the integration of information from different sources to form a reliable basis for behavior. While animals behave as if they optimally integrate information according to Bayesian probability theory, the implementation of the required computations in the biological substrate remains unclear. We propose a novel, Bayesian view on the dynamics of conductance-based neurons and synapses which suggests that they are naturally equipped to optimally perform information integration. In our approach apical dendrites represent prior expectations over somatic potentials, while basal dendrites represent likelihoods of somatic potentials. These are parametrized by local quantities, the effective reversal potentials and membrane conductances. We formally demonstrate that under these assumptions the somatic compartment naturally computes the corresponding posterior. We derive a gradient-based plasticity rule, allowing neurons to learn desired target distributions and weight synaptic inputs by their relative reliabilities. Our theory explains various experimental findings on the system and single-cell level related to multi-sensory integration, which we illustrate with simulations. Furthermore, we make experimentally testable predictions on Bayesian dendritic integration and synaptic plasticity.

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arXiv2022

Versatile emulation of spiking neural networks on an accelerated neuromorphic substrate

We present first experimental results on the novel BrainScaleS-2 neuromorphic architecture based on an analog neuro-synaptic core and augmented by embedded microprocessors for complex plasticity and experiment control. The high acceleration factor of 1000 compared to biological dynamics enables the execution of computationally expensive tasks, by allowing the fast emulation of long-duration experiments or rapid iteration over many consecutive trials. The flexibility of our architecture is demonstrated in a suite of five distinct experiments, which emphasize different aspects of the BrainScaleS-2 system.

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arXiv2022

Cortical oscillations implement a backbone for sampling-based computation in spiking neural networks

Being permanently confronted with an uncertain world, brains have faced evolutionary pressure to represent this uncertainty in order to respond appropriately. Often, this requires visiting multiple interpretations of the available information or multiple solutions to an encountered problem. This gives rise to the so-called mixing problem: since all of these "valid" states represent powerful attractors, but between themselves can be very dissimilar, switching between such states can be difficult. We propose that cortical oscillations can be effectively used to overcome this challenge. By acting as an effective temperature, background spiking activity modulates exploration. Rhythmic changes induced by cortical oscillations can then be interpreted as a form of simulated tempering. We provide a rigorous mathematical discussion of this link and study some of its phenomenological implications in computer simulations. This identifies a new computational role of cortical oscillations and connects them to various phenomena in the brain, such as sampling-based probabilistic inference, memory replay, multisensory cue combination, and place cell flickering.

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arXiv2022

Learning cortical representations through perturbed and adversarial dreaming

Humans and other animals learn to extract general concepts from sensory experience without extensive teaching. This ability is thought to be facilitated by offline states like sleep where previous experiences are systemically replayed. However, the characteristic creative nature of dreams suggests that learning semantic representations may go beyond merely replaying previous experiences. We support this hypothesis by implementing a cortical architecture inspired by generative adversarial networks (GANs). Learning in our model is organized across three different global brain states mimicking wakefulness, NREM and REM sleep, optimizing different, but complementary objective functions. We train the model on standard datasets of natural images and evaluate the quality of the learned representations. Our results suggest that generating new, virtual sensory inputs via adversarial dreaming during REM sleep is essential for extracting semantic concepts, while replaying episodic memories via perturbed dreaming during NREM sleep improves the robustness of latent representations. The model provides a new computational perspective on sleep states, memory replay and dreams and suggests a cortical implementation of GANs.

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arXiv2021

Latent Equilibrium: A unified learning theory for arbitrarily fast computation with arbitrarily slow neurons

The response time of physical computational elements is finite, and neurons are no exception. In hierarchical models of cortical networks each layer thus introduces a response lag. This inherent property of physical dynamical systems results in delayed processing of stimuli and causes a timing mismatch between network output and instructive signals, thus afflicting not only inference, but also learning. We introduce Latent Equilibrium, a new framework for inference and learning in networks of slow components which avoids these issues by harnessing the ability of biological neurons to phase-advance their output with respect to their membrane potential. This principle enables quasi-instantaneous inference independent of network depth and avoids the need for phased plasticity or computationally expensive network relaxation phases. We jointly derive disentangled neuron and synapse dynamics from a prospective energy function that depends on a network's generalized position and momentum. The resulting model can be interpreted as a biologically plausible approximation of error backpropagation in deep cortical networks with continuous-time, leaky neuronal dynamics and continuously active, local plasticity. We demonstrate successful learning of standard benchmark datasets, achieving competitive performance using both fully-connected and convolutional architectures, and show how our principle can be applied to detailed models of cortical microcircuitry. Furthermore, we study the robustness of our model to spatio-temporal substrate imperfections to demonstrate its feasibility for physical realization, be it in vivo or in silico.

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arXiv2021

Fast and energy-efficient neuromorphic deep learning with first-spike times

For a biological agent operating under environmental pressure, energy consumption and reaction times are of critical importance. Similarly, engineered systems are optimized for short time-to-solution and low energy-to-solution characteristics. At the level of neuronal implementation, this implies achieving the desired results with as few and as early spikes as possible. With time-to-first-spike coding both of these goals are inherently emerging features of learning. Here, we describe a rigorous derivation of a learning rule for such first-spike times in networks of leaky integrate-and-fire neurons, relying solely on input and output spike times, and show how this mechanism can implement error backpropagation in hierarchical spiking networks. Furthermore, we emulate our framework on the BrainScaleS-2 neuromorphic system and demonstrate its capability of harnessing the system's speed and energy characteristics. Finally, we examine how our approach generalizes to other neuromorphic platforms by studying how its performance is affected by typical distortive effects induced by neuromorphic substrates.

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arXiv2021

Evolving Neuronal Plasticity Rules using Cartesian Genetic Programming

We formulate the search for phenomenological models of synaptic plasticity as an optimization problem. We employ Cartesian genetic programming to evolve biologically plausible human-interpretable plasticity rules that allow a given network to successfully solve tasks from specific task families. While our evolving-to-learn approach can be applied to various learning paradigms, here we illustrate its power by evolving plasticity rules that allow a network to efficiently determine the first principal component of its input distribution. We demonstrate that the evolved rules perform competitively with known hand-designed solutions. We explore how the statistical properties of the datasets used during the evolutionary search influences the form of the plasticity rules and discover new rules which are adapted to the structure of the corresponding datasets.

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