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Eric Shea-Brown

Publications and source records attributed to Eric Shea-Brown.

At least 19 recordsLinked to original sources
arXiv2026

Center-Manifold Reduction of Learning at Bifurcations: Interference and Rich Learning in Recurrent Neural Networks

Rich learning in recurrent neural networks often proceeds through sudden transitions in latent dynamics, but there is little theory predicting how gradient descent behaves during these events. We study the local learning geometry near codimension-one bifurcations through the global empirical Neural Tangent Kernel (GeNTK). Under local center-manifold conditions, and when bifurcation-related sensitivity dominates bounded residual terms, we show that the global parameter-to-state Jacobian \(D_\theta h\) is approximated by a low-rank normal-form operator. The induced GeNTK and Fisher information matrix therefore become strongly amplified and anisotropic, concentrating toward a rank-one channel for the four scalar codimension-one bifurcations and a rank-two real channel for a Neimark--Sacker bifurcation. Controlled high-dimensional RNN experiments validate this operator reduction. In learned RNNs, the same low-rank concentration coincides with abrupt loss changes and subtask interference, while a local projection predicts the sign of these effects near isolated events. Finally, in an input-driven 15-task LeakyRNN, GeNTK amplification aligns with continuation-detected changes in the MemoryPro dynamics. These results suggest a tractable operator-level description of learning near dynamical transitions, together with scalable diagnostics for amplified low-dimensional learning geometry.

cs.LG↗
arXiv2026

The Global Empirical NTK: Self-Referential Bias and Dimensionality of Gradient Descent Learning

In training a neural network with gradient descent (GD), each iteration induces a linear operator that governs first-order updates to a model's internal state variables. We define this operator as the Global Empirical Neural Tangent Kernel (NTK). In finite-width networks, the NTK is typically intractable to form, leading prior work to focus on restrictive settings such as tracking outputs only or taking infinite-width limits. Here, we study the structure of the NTK for a range of models. Formulating the model state as the solution to a single global implicit constraint, we derive the NTK as a product of two operators: K, accounting for immediate parameter-to-state interactions, and P, describing internal state-to-state dependencies. For a broad class of weight-based models, including RNNs and transformers, we prove a universal Kronecker-core theorem showing that K admits an exact, computable form given by the Gram matrix of weight-site variables. This core structure reveals that the NTK is structurally bottlenecked, constraining its effective rank and giving rise to a self-referential bias whereby GD preferentially learns within dominant modes of joint hidden and input activity. For recurrent models, we examine the spectrum of the NTK and show when it is biased and low-rank in space or time under the proposed decomposition. We further demonstrate that model dynamics at initialization bias the NTK, restricting learning and preventing task components from being learned effectively. Finally, we show that the NTK associated with a self-attention transformer is likewise structurally constrained to be low-rank. Overall, we show that the NTK possesses tractable structure that explains GD bias toward task solutions and the emergence of low-rank representations. To enable use of the NTK as a practical metric, we build kpflow, a library relying on randomized matrix-free numerical linear algebra.

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arXiv2026

Identifying Connectivity Distributions from Neural Dynamics Using Flows

Connectivity structure shapes neural computation, but inferring this structure from population recordings is degenerate: multiple connectivity structures can generate identical dynamics. Recent work uses low-rank recurrent neural networks (lrRNNs) to infer low-dimensional latent dynamics and connectivity from observed activity, enabling a mechanistic interpretation of the dynamics. However, standard approaches for training lrRNNs can recover spurious structures irrelevant to the underlying dynamics. We first characterize the identifiability of connectivity structures in lrRNNs and determine conditions under which a unique solution exists. To find such solutions, we develop an inference framework based on maximum entropy and continuous normalizing flows (CNFs), trained via flow matching. Instead of estimating a single connectivity matrix, our method learns a distribution over connection weights that is maximally unbiased over unidentifiable components while matching the observed dynamics. This approach captures complex yet necessary distributions such as heavy-tailed connectivity found in empirical data. We validate our method on synthetic datasets with connectivity structures that generate multistable attractors, limit cycles, and ring attractors, and demonstrate its applicability in recordings from rat frontal cortex during decision-making. Our framework shifts circuit inference from recovering connectivity to identifying which connectivity structures are computationally required, and which are artifacts of underconstrained inference.

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arXiv2025

Benchmarking Probabilistic Time Series Forecasting Models on Neural Activity

Neural activity forecasting is central to understanding neural systems and enabling closed-loop control. While deep learning has recently advanced the state-of-the-art in the time series forecasting literature, its application to neural activity forecasting remains limited. To bridge this gap, we systematically evaluated eight probabilistic deep learning models, including two foundation models, that have demonstrated strong performance on general forecasting benchmarks. We compared them against four classical statistical models and two baseline methods on spontaneous neural activity recorded from mouse cortex via widefield imaging. Across prediction horizons, several deep learning models consistently outperformed classical approaches, with the best model producing informative forecasts up to 1.5 seconds into the future. Our findings point toward future control applications and open new avenues for probing the intrinsic temporal structure of neural activity.

cs.LG↗
arXiv2025

Interpretable time series analysis with Gumbel dynamics

Switching dynamical systems can model complicated time series data while maintaining interpretability by inferring a finite set of dynamics primitives and explaining different portions of the observed time series with one of these primitives. However, due to the discrete nature of this set, such models struggle to capture smooth, variable-speed transitions, as well as stochastic mixtures of overlapping states, and the inferred dynamics often display spurious rapid switching on real-world datasets. Here, we propose the Gumbel Dynamical Model (GDM). First, by introducing a continuous relaxation of discrete states and a different noise model defined on the relaxed-discrete state space via the Gumbel distribution, GDM expands the set of available state dynamics, allowing the model to approximate smoother and non-stationary ground-truth dynamics more faithfully. Second, the relaxation makes the model fully differentiable, enabling fast and scalable training with standard gradient descent methods. We validate our approach on standard simulation datasets and highlight its ability to model soft, sticky states and transitions in a stochastic setting. Furthermore, we apply our model to two real-world datasets, demonstrating its ability to infer interpretable states in stochastic time series with multiple dynamics, a setting where traditional methods often fail.

cs.LG↗
arXiv2025

KPFlow: An Operator Perspective on Dynamic Collapse Under Gradient Descent Training of Recurrent Networks

Gradient Descent (GD) and its variants are the primary tool for enabling efficient training of recurrent dynamical systems such as Recurrent Neural Networks (RNNs), Neural ODEs and Gated Recurrent units (GRUs). The dynamics that are formed in these models exhibit features such as neural collapse and emergence of latent representations that may support the remarkable generalization properties of networks. In neuroscience, qualitative features of these representations are used to compare learning in biological and artificial systems. Despite recent progress, there remains a need for theoretical tools to rigorously understand the mechanisms shaping learned representations, especially in finite, non-linear models. Here, we show that the gradient flow, which describes how the model's dynamics evolve over GD, can be decomposed into a product that involves two operators: a Parameter Operator, K, and a Linearized Flow Propagator, P. K mirrors the Neural Tangent Kernel in feed-forward neural networks, while P appears in Lyapunov stability and optimal control theory. We demonstrate two applications of our decomposition. First, we show how their interplay gives rise to low-dimensional latent dynamics under GD, and, specifically, how the collapse is a result of the network structure, over and above the nature of the underlying task. Second, for multi-task training, we show that the operators can be used to measure how objectives relevant to individual sub-tasks align. We experimentally and theoretically validate these findings, providing an efficient Pytorch package, \emph{KPFlow}, implementing robust analysis tools for general recurrent architectures. Taken together, our work moves towards building a next stage of understanding of GD learning in non-linear recurrent models.

cs.LG↗
arXiv2024

Identifying the impact of local connectivity patterns on dynamics in excitatory-inhibitory networks

Networks of excitatory and inhibitory (EI) neurons form a canonical circuit in the brain. Seminal theoretical results on dynamics of such networks are based on the assumption that synaptic strengths depend on the type of neurons they connect, but are otherwise statistically independent. Recent synaptic physiology datasets however highlight the prominence of specific connectivity patterns that go well beyond what is expected from independent connections. While decades of influential research have demonstrated the strong role of the basic EI cell type structure, to which extent additional connectivity features influence dynamics remains to be fully determined. Here we examine the effects of pairwise connectivity motifs on the linear dynamics in EI networks using an analytical framework that approximates the connectivity in terms of low-rank structures. This low-rank approximation is based on a mathematical derivation of the dominant eigenvalues of the connectivity matrix and predicts the impact on responses to external inputs of connectivity motifs and their interactions with cell-type structure. Our results reveal that a particular pattern of connectivity, chain motifs, have a much stronger impact on dominant eigenmodes than other pairwise motifs. An overrepresentation of chain motifs induces a strong positive eigenvalue in inhibition-dominated networks and generates a potential instability that requires revisiting the classical excitation-inhibition balance criteria. Examining effects of external inputs, we show that chain motifs can on their own induce paradoxical responses where an increased input to inhibitory neurons leads to a decrease in their activity due to the recurrent feedback. These findings have direct implications for the interpretation of experiments in which responses to optogenetic perturbations are measured and used to infer the dynamical regime of cortical circuits.

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arXiv2023

Evolutionary algorithms as an alternative to backpropagation for supervised training of Biophysical Neural Networks and Neural ODEs

Training networks consisting of biophysically accurate neuron models could allow for new insights into how brain circuits can organize and solve tasks. We begin by analyzing the extent to which the central algorithm for neural network learning -- stochastic gradient descent through backpropagation (BP) -- can be used to train such networks. We find that properties of biophysically based neural network models needed for accurate modelling such as stiffness, high nonlinearity and long evaluation timeframes relative to spike times makes BP unstable and divergent in a variety of cases. To address these instabilities and inspired by recent work, we investigate the use of "gradient-estimating" evolutionary algorithms (EAs) for training biophysically based neural networks. We find that EAs have several advantages making them desirable over direct BP, including being forward-pass only, robust to noisy and rigid losses, allowing for discrete loss formulations, and potentially facilitating a more global exploration of parameters. We apply our method to train a recurrent network of Morris-Lecar neuron models on a stimulus integration and working memory task, and show how it can succeed in cases where direct BP is inapplicable. To expand on the viability of EAs in general, we apply them to a general neural ODE problem and a stiff neural ODE benchmark and find again that EAs can out-perform direct BP here, especially for the over-parameterized regime. Our findings suggest that biophysical neurons could provide useful benchmarks for testing the limits of BP-adjacent methods, and demonstrate the viability of EAs for training networks with complex components.

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arXiv2023

Expressive probabilistic sampling in recurrent neural networks

In sampling-based Bayesian models of brain function, neural activities are assumed to be samples from probability distributions that the brain uses for probabilistic computation. However, a comprehensive understanding of how mechanistic models of neural dynamics can sample from arbitrary distributions is still lacking. We use tools from functional analysis and stochastic differential equations to explore the minimum architectural requirements for $\textit{recurrent}$ neural circuits to sample from complex distributions. We first consider the traditional sampling model consisting of a network of neurons whose outputs directly represent the samples (sampler-only network). We argue that synaptic current and firing-rate dynamics in the traditional model have limited capacity to sample from a complex probability distribution. We show that the firing rate dynamics of a recurrent neural circuit with a separate set of output units can sample from an arbitrary probability distribution. We call such circuits reservoir-sampler networks (RSNs). We propose an efficient training procedure based on denoising score matching that finds recurrent and output weights such that the RSN implements Langevin sampling. We empirically demonstrate our model's ability to sample from several complex data distributions using the proposed neural dynamics and discuss its applicability to developing the next generation of sampling-based brain models.

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arXiv2023

Attention for Causal Relationship Discovery from Biological Neural Dynamics

This paper explores the potential of the transformer models for learning Granger causality in networks with complex nonlinear dynamics at every node, as in neurobiological and biophysical networks. Our study primarily focuses on a proof-of-concept investigation based on simulated neural dynamics, for which the ground-truth causality is known through the underlying connectivity matrix. For transformer models trained to forecast neuronal population dynamics, we show that the cross attention module effectively captures the causal relationship among neurons, with an accuracy equal or superior to that for the most popular Granger causality analysis method. While we acknowledge that real-world neurobiology data will bring further challenges, including dynamic connectivity and unobserved variability, this research offers an encouraging preliminary glimpse into the utility of the transformer model for causal representation learning in neuroscience.

cs.LG↗
arXiv2023

How connectivity structure shapes rich and lazy learning in neural circuits

In theoretical neuroscience, recent work leverages deep learning tools to explore how some network attributes critically influence its learning dynamics. Notably, initial weight distributions with small (resp. large) variance may yield a rich (resp. lazy) regime, where significant (resp. minor) changes to network states and representation are observed over the course of learning. However, in biology, neural circuit connectivity could exhibit a low-rank structure and therefore differs markedly from the random initializations generally used for these studies. As such, here we investigate how the structure of the initial weights -- in particular their effective rank -- influences the network learning regime. Through both empirical and theoretical analyses, we discover that high-rank initializations typically yield smaller network changes indicative of lazier learning, a finding we also confirm with experimentally-driven initial connectivity in recurrent neural networks. Conversely, low-rank initialization biases learning towards richer learning. Importantly, however, as an exception to this rule, we find lazier learning can still occur with a low-rank initialization that aligns with task and data statistics. Our research highlights the pivotal role of initial weight structures in shaping learning regimes, with implications for metabolic costs of plasticity and risks of catastrophic forgetting.

cs.NE↗
arXiv2023

A simple connection from loss flatness to compressed neural representations

Despite extensive study, the significance of sharpness -- the trace of the loss Hessian at local minima -- remains unclear. We investigate an alternative perspective: how sharpness relates to the geometric structure of neural representations, specifically representation compression, defined as how strongly neural activations concentrate under local input perturbations. We introduce three measures -- Local Volumetric Ratio (LVR), Maximum Local Sensitivity (MLS), and Local Dimensionality -- and derive upper bounds showing these are mathematically constrained by sharpness: flatter minima necessarily limit compression. We extend these bounds to reparametrization-invariant sharpness and introduce network-wide variants (NMLS, NVR) that provide tighter, more stable bounds than prior single-layer analyses. Empirically, we validate consistent positive correlations across feedforward, convolutional, and transformer architectures. Our results suggest that sharpness fundamentally quantifies representation compression, offering a principled resolution to contradictory findings on the sharpness-generalization relationship.

cs.LG↗
arXiv2023

Beyond accuracy: generalization properties of bio-plausible temporal credit assignment rules

To unveil how the brain learns, ongoing work seeks biologically-plausible approximations of gradient descent algorithms for training recurrent neural networks (RNNs). Yet, beyond task accuracy, it is unclear if such learning rules converge to solutions that exhibit different levels of generalization than their nonbiologically-plausible counterparts. Leveraging results from deep learning theory based on loss landscape curvature, we ask: how do biologically-plausible gradient approximations affect generalization? We first demonstrate that state-of-the-art biologically-plausible learning rules for training RNNs exhibit worse and more variable generalization performance compared to their machine learning counterparts that follow the true gradient more closely. Next, we verify that such generalization performance is correlated significantly with loss landscape curvature, and we show that biologically-plausible learning rules tend to approach high-curvature regions in synaptic weight space. Using tools from dynamical systems, we derive theoretical arguments and present a theorem explaining this phenomenon. This predicts our numerical results, and explains why biologically-plausible rules lead to worse and more variable generalization properties. Finally, we suggest potential remedies that could be used by the brain to mitigate this effect. To our knowledge, our analysis is the first to identify the reason for this generalization gap between artificial and biologically-plausible learning rules, which can help guide future investigations into how the brain learns solutions that generalize.

cs.NE↗
arXiv2023

Biologically-plausible backpropagation through arbitrary timespans via local neuromodulators

The spectacular successes of recurrent neural network models where key parameters are adjusted via backpropagation-based gradient descent have inspired much thought as to how biological neuronal networks might solve the corresponding synaptic credit assignment problem. There is so far little agreement, however, as to how biological networks could implement the necessary backpropagation through time, given widely recognized constraints of biological synaptic network signaling architectures. Here, we propose that extra-synaptic diffusion of local neuromodulators such as neuropeptides may afford an effective mode of backpropagation lying within the bounds of biological plausibility. Going beyond existing temporal truncation-based gradient approximations, our approximate gradient-based update rule, ModProp, propagates credit information through arbitrary time steps. ModProp suggests that modulatory signals can act on receiving cells by convolving their eligibility traces via causal, time-invariant and synapse-type-specific filter taps. Our mathematical analysis of ModProp learning, together with simulation results on benchmark temporal tasks, demonstrate the advantage of ModProp over existing biologically-plausible temporal credit assignment rules. These results suggest a potential neuronal mechanism for signaling credit information related to recurrent interactions over a longer time horizon. Finally, we derive an in-silico implementation of ModProp that could serve as a low-complexity and causal alternative to backpropagation through time.

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arXiv2020

Network Dynamics Governed by Lyapunov Functions: From Memory to Classification

In 1982 John Hopfield published a neural network model for memory retrieval, a model that became a cornerstone in theoretical neuroscience. A key ingredient of the Hopfield model was the use of a network dynamics that is governed by a Lyapunov function. In a recent paper, Krotov and Hopfield showed how a Lyapunov function governs a biological plausible learning rule for the neural networks' connectivity. By doing so, they bring an intriguing approach to classification tasks, and show the relevance of the broader framework across decades in the field.

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arXiv2020

Linking structure and activity in nonlinear spiking networks

Recent experimental advances are producing an avalanche of data on both neural connectivity and neural activity. To take full advantage of these two emerging datasets we need a framework that links them, revealing how collective neural activity arises from the structure of neural connectivity and intrinsic neural dynamics. This problem of {\it structure-driven activity} has drawn major interest in computational neuroscience. Existing methods for relating activity and architecture in spiking networks rely on linearizing activity around a central operating point and thus fail to capture the nonlinear responses of individual neurons that are the hallmark of neural information processing. Here, we overcome this limitation and present a new relationship between connectivity and activity in networks of nonlinear spiking neurons by developing a diagrammatic fluctuation expansion based on statistical field theory. We explicitly show how recurrent network structure produces pairwise and higher-order correlated activity, and how nonlinearities impact the networks' spiking activity. Our findings open new avenues to investigating how single-neuron nonlinearities---including those of different cell types---combine with connectivity to shape population activity and function.

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arXiv2019

Comparison Against Task Driven Artificial Neural Networks Reveals Functional Organization of Mouse Visual Cortex

Partially inspired by features of computation in visual cortex, deep neural networks compute hierarchical representations of their inputs. While these networks have been highly successful in machine learning, it remains unclear to what extent they can aid our understanding of cortical function. Several groups have developed metrics that provide a quantitative comparison between representations computed by networks and representations measured in cortex. At the same time, neuroscience is well into an unprecedented phase of large-scale data collection, as evidenced by projects such as the Allen Brain Observatory. Despite the magnitude of these efforts, in a given experiment only a fraction of units are recorded, limiting the information available about the cortical representation. Moreover, only a finite number of stimuli can be shown to an animal over the course of a realistic experiment. These limitations raise the question of how and whether metrics that compare representations of deep networks are meaningful on these datasets. Here, we empirically quantify the capabilities and limitations of these metrics due to limited image presentations and neuron samples. We find that the comparison procedure is robust to different choices of stimuli set and the level of subsampling that one might expect in a large-scale brain survey with thousands of neurons. Using these results, we compare the representations measured in the Allen Brain Observatory in response to natural image presentations to deep neural network. We show that the visual cortical areas are relatively high order representations (in that they map to deeper layers of convolutional neural networks). Furthermore, we see evidence of a broad, more parallel organization rather than a sequential hierarchy, with the primary area VISp(V1) being lower order relative to the other areas.

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