arXiv subjects

Steven W. Zucker

Publications and source records attributed to Steven W. Zucker.

At least 19 recordsLinked to original sources

arXiv2026

Bracketing Uncertainty in Clustering Under the Manifold Hypothesis

The manifold hypothesis suggests a natural criterion for clustering: partition data according to the manifold component from which each point is drawn. Whether two components are separable depends on a geometric tradeoff: the ambient separation between components versus the largest gap in sampling. In practice, this tradeoff is rarely assessed explicitly, leading standard methods to over-commit to a single clustering assignment even when the data do not support a unique answer. We formalize this tradeoff by combining intrinsic manifold geometry (volume growth and reach) with sample-level quantities (fill distance and density), yielding a threshold phenomenon for mutual-$k$-nearest-neighbor graphs: when the offset-to-fill ratio exceeds a conservative upper threshold, component separation is preserved; below a lower threshold, components fuse. The gap between these thresholds defines a geometric uncertainty zone in which the number of clusters is not identifiable from the data. Nevertheless, conventional approaches still seek one: sweeping parameters (an engineering approach) or fitting a generative mixture model (a model-based approach). Rather than forcing a single estimate of the number of clusters, we propose Manifold-Based Clustering (MBC), which returns an explicit bracket interval to quantify the underlying data uncertainty. This bracket acts as an empirically calibrated diagnostic: it narrows when a single resolution is supported, widens when multiple resolutions coexist, and collapses to one when no separated structure is detectable. Empirically, we find that many real datasets lie within the uncertainty zone rather than admitting one clear answer. Our results suggest that ambiguity in cluster number is often intrinsic, and should be quantified rather than resolved.

stat.ML↗

arXiv2026

Inferring Active Neural Circuits Using Diffusion Scores

In biological systems, neural circuits compute through directed, short-latency interactions whose effects unfold across multiple time scales and behavioral contexts. We address the problem of inferring these local, lag-specific interactions from sampled neural population activity under varying stimuli, without assuming a parametric form for the underlying dynamics. Our approach leverages denoising score models by estimating joint-window scores over consecutive activity snapshots (i.e., brain states) and converting these scores into calibrated, directed edge tests via cross-block score products. The key insight is that these products recover the Jacobian of the transition map between brain states under nonlinear dynamics. To cleanly separate lag-specific effects, we introduce minimal multi-block windows that condition on intermediate time points, avoiding the omitted-lag bias inherent in pairwise analyses. The resulting method, Score--Block Time Graphs (SBTG), identifies lag-specific directed interactions in sampled neuronal population data. We specifically apply SBTG to whole-brain C. elegans calcium imaging data to recover lag-specific circuit structure not resolved by current methods, including improved alignment with independent connectomes, cell-type-specific temporal organization, and neuromodulatory profiles consistent with known receptor kinetics. These findings highlight the potential for SBTG to serve as a practical ``AI for science'' tool by turning high-dimensional neural population recordings into statistically testable circuit hypotheses.

q-bio.NC↗

arXiv2026

Decoding Alignment without Encoding Alignment: A critique of similarity analysis in neuroscience

Decoding approaches are widely used in neuroscience and machine learning to compare stimulus representations across neural systems, such as different brain regions, organisms, and deep learning models. Popular methods include decoding (perceptual) manifolds and alignment metrics such as Representational Similarity Analysis (RSA) and Dynamic Similarity Analysis (DSA), where similarity in decoding representations is interpreted as evidence for similar computation. This paper demonstrates a fundamental weakness behind this approach: it is misleading to assume that representational geometry is representative of a neuronal population as a whole, when such representations may actually be shaped by a very small subset of neurons. We show that the complementary encoding paradigm addresses this issue directly: it characterizes how neurons are organized globally in terms of their responses to a set of data, providing insight into how the decoding representation is implemented by neurons within a population. We demonstrate across experiments in biological systems and deep learning models that (i) surprisingly, similar decoding behavior and high representational alignment can arise from small, non-representative subpopulations of neurons; and critically, (ii) alignment metrics are insensitive to encoding manifold topology (how function is distributed across neurons), despite this being a key signature of differentiation across biological systems. A controlled MNIST experiment provides causal evidence: decoding metrics remain unchanged even when encoding topology is causally manipulated via the training loss. Overall, similarity in decoding behavior, as measured by classic alignment metrics, does not imply similarity in function or computation, motivating the use of encoding manifolds as a complementary tool for comparing neural systems.

q-bio.NC↗

arXiv2026

NeuroAI and Beyond: Bridging Between Advances in Neuroscience and ArtificialIntelligence

Neuroscience and Artificial Intelligence (AI) have made impressive progress in recent years but remain only loosely interconnected. Based on a workshop convened by the National Science Foundation in August 2025, we identify three fundamental capability gaps in current AI: the inability to interact with the physical world, inadequate learning that produces brittle systems, and unsustainable energy and data inefficiency. We describe the neuroscience principles that address each: co-design of body and controller, prediction through interaction, multi-scale learning with neuromodulatory control, hierarchical distributed architectures, and sparse event-driven computation. We present a research roadmap organized around these principles at near, mid, and long-term horizons. We argue that realizing this program requires a new generation of researchers trained across the boundary between neuroscience and engineering, and describe the institutional conditions: interdisciplinary training, hardware access, community standards, and ethics, needed to support them. We conclude that NeuroAI, neuroscience-informed artificial intelligence, has the potential to overcome limitations of current AI while deepening our understanding of biological neural computation.

q-bio.NC↗

arXiv2026

How 'Neural' is a Neural Foundation Model?

Foundation models have shown remarkable success in fitting biological visual systems; however, their black-box nature inherently limits their utility for understanding brain function. Here, we peek inside a SOTA foundation model of neural activity (Wang et al., 2025) as a physiologist might, characterizing each 'neuron' based on its temporal response properties to parametric stimuli. We analyze how different stimuli are represented in neural activity space by building decoding manifolds, and we analyze how different neurons are represented in stimulus-response space by building neural encoding manifolds. We find that the different processing stages of the model (i.e., the feedforward encoder, recurrent, and readout modules) each exhibit qualitatively different representational structures in these manifolds. The recurrent module shows a jump in capabilities over the encoder module by 'pushing apart' the representations of different temporal stimulus patterns. Our 'tubularity' metric quantifies this stimulus-dependent development of neural activity as biologically plausible. The readout module achieves high fidelity by using numerous specialized feature maps rather than biologically plausible mechanisms. Overall, this study provides a window into the inner workings of a prominent neural foundation model, gaining insights into the biological relevance of its internals through the novel analysis of its neurons' joint temporal response patterns. Our findings suggest design changes that could bring neural foundation models into closer alignment with biological systems: introducing recurrence in early encoder stages, and constraining features in the readout module.

q-bio.NC↗

arXiv2025

Manifolds and Modules: How Function Develops in a Neural Foundation Model

Foundation models have shown remarkable success in fitting biological visual systems; however, their black-box nature inherently limits their utility for understanding brain function. Here, we peek inside a SOTA foundation model of neural activity (Wang et al., 2025) as a physiologist might, characterizing each 'neuron' based on its temporal response properties to parametric stimuli. We analyze how different stimuli are represented in neural activity space by building decoding manifolds, and we analyze how different neurons are represented in stimulus-response space by building neural encoding manifolds. We find that the different processing stages of the model (i.e., the feedforward encoder, recurrent, and readout modules) each exhibit qualitatively different representational structures in these manifolds. The recurrent module shows a jump in capabilities over the encoder module by 'pushing apart' the representations of different temporal stimulus patterns; while the readout module achieves biological fidelity by using numerous specialized feature maps rather than biologically plausible mechanisms. Overall, we present this work as a study of the inner workings of a prominent neural foundation model, gaining insights into the biological relevance of its internals through the novel analysis of its neurons' joint temporal response patterns.

q-bio.NC↗

arXiv2024

A separability-based approach to quantifying generalization: which layer is best?

Generalization to unseen data remains poorly understood for deep learning classification and foundation models, especially in the open set scenario. How can one assess the ability of networks to adapt to new or extended versions of their input space in the spirit of few-shot learning, out-of-distribution generalization, domain adaptation, and category discovery? Which layers of a network are likely to generalize best? We provide a new method for evaluating the capacity of networks to represent a sampled domain, regardless of whether the network has been trained on all classes in that domain. Our approach is the following: after fine-tuning state-of-the-art pre-trained models for visual classification on a particular domain, we assess their performance on data from related but distinct variations in that domain. Generalization power is quantified as a function of the latent embeddings of unseen data from intermediate layers for both unsupervised and supervised settings. Working throughout all stages of the network, we find that (i) high classification accuracy does not imply high generalizability; and (ii) deeper layers in a model do not always generalize the best, which has implications for pruning. Since the trends observed across datasets are largely consistent, we conclude that our approach reveals (a function of) the intrinsic capacity of the different layers of a model to generalize. Our code is available at https://github.com/dyballa/generalization

cs.LG↗

arXiv2024

Individuation of 3D perceptual units from neurogeometry of binocular cells

We model the functional architecture of the early stages of three-dimensional vision by extending the neurogeometric sub-Riemannian model for stereo-vision introduced in \cite{BCSZ23}. A new framework for correspondence is introduced that integrates a neural-based algorithm to achieve stereo correspondence locally while, simultaneously, organizing the corresponding points into global perceptual units. The result is an effective scene segmentation. We achieve this using harmonic analysis on the sub-Riemannian structure and show, in a comparison against Riemannian distance, that the sub-Riemannian metric is central to the solution.

q-bio.NC↗

arXiv2024

Zero-shot generalization across architectures for visual classification

Generalization to unseen data is a key desideratum for deep networks, but its relation to classification accuracy is unclear. Using a minimalist vision dataset and a measure of generalizability, we show that popular networks, from deep convolutional networks (CNNs) to transformers, vary in their power to extrapolate to unseen classes both across layers and across architectures. Accuracy is not a good predictor of generalizability, and generalization varies non-monotonically with layer depth.

cs.CV↗

arXiv2024

Learning dynamic representations of the functional connectome in neurobiological networks

The static synaptic connectivity of neuronal circuits stands in direct contrast to the dynamics of their function. As in changing community interactions, different neurons can participate actively in various combinations to effect behaviors at different times. We introduce an unsupervised approach to learn the dynamic affinities between neurons in live, behaving animals, and to reveal which communities form among neurons at different times. The inference occurs in two major steps. First, pairwise non-linear affinities between neuronal traces from brain-wide calcium activity are organized by non-negative tensor factorization (NTF). Each factor specifies which groups of neurons are most likely interacting for an inferred interval in time, and for which animals. Finally, a generative model that allows for weighted community detection is applied to the functional motifs produced by NTF to reveal a dynamic functional connectome. Since time codes the different experimental variables (e.g., application of chemical stimuli), this provides an atlas of neural motifs active during separate stages of an experiment (e.g., stimulus application or spontaneous behaviors). Results from our analysis are experimentally validated, confirming that our method is able to robustly predict causal interactions between neurons to generate behavior. Code is available at https://github.com/dyballa/dynamic-connectomes.

q-bio.NC↗

arXiv2023

Good continuation in 3D: the neurogeometry of stereo vision

Classical good continuation for image curves is based on $2D$ position and orientation. It is supported by the columnar organization of cortex, by psychophysical experiments, and by rich models of (differential) geometry. Here we extend good continuation to stereo. We introduce a neurogeometric model, in which the parametrizations involve both spatial and orientation disparities. Our model provides insight into the neurobiology, suggesting an implicit organization for neural interactions and a well-defined $3D$ association field. Our model sheds light on the computations underlying the correspondence problem, and illustrates how good continuation in the world generalizes good continuation in the plane.

q-bio.NC↗

arXiv2023

IAN: Iterated Adaptive Neighborhoods for manifold learning and dimensionality estimation

Invoking the manifold assumption in machine learning requires knowledge of the manifold's geometry and dimension, and theory dictates how many samples are required. However, in applications data are limited, sampling may not be uniform, and manifold properties are unknown and (possibly) non-pure; this implies that neighborhoods must adapt to the local structure. We introduce an algorithm for inferring adaptive neighborhoods for data given by a similarity kernel. Starting with a locally-conservative neighborhood (Gabriel) graph, we sparsify it iteratively according to a weighted counterpart. In each step, a linear program yields minimal neighborhoods globally and a volumetric statistic reveals neighbor outliers likely to violate manifold geometry. We apply our adaptive neighborhoods to non-linear dimensionality reduction, geodesic computation and dimension estimation. A comparison against standard algorithms using, e.g., k-nearest neighbors, demonstrates their usefulness. Code for our algorithm will be available at https://github.com/dyballa/IAN

cs.LG↗

arXiv2020

From Boundaries to Bumps: when closed (extremal) contours are critical

Invariants underlying shape inference are elusive: a variety of shapes can give rise to the same image, and a variety of images can be rendered from the same shape. The occluding contour is a rare exception: it has both image salience, in terms of isophotes, and surface meaning, in terms of surface normal. We relax the notion of occluding contour to define closed extremal curves, a new shape invariant that exists at the topological level. They surround bumps, a common but ill-specified interior shape component, and formalize the qualitative nature of bump perception. Extremal curves are biologically computable, unify shape inferences from shading, texture, and specular materials, and predict new phenomena in bump perception.

cs.CV↗

arXiv2018

Tensors, Differential Geometry and Statistical Shading Analysis

We develop a linear algebraic framework for the shape-from-shading problem, because tensors arise when scalar (e.g. image) and vector (e.g. surface normal) fields are differentiated multiple times. Using this framework, we first investigate when image derivatives exhibit invariance to changing illumination by calculating the statistics of image derivatives under general distributions on the light source. Second, we apply that framework to develop Taylor-like expansions, and build a boot-strapping algorithm to find the polynomial surface solutions (under any light source) consistent with a given patch to arbitrary order. A generic constraint on the light source restricts these solutions to a 2-D subspace, plus an unknown rotation matrix. It is this unknown matrix that encapsulates the ambiguity in the problem. Finally, we use the framework to computationally validate the hypothesis that image orientations (derivatives) provide increased invariance to illumination by showing (for a Lambertian model) that a shape-from-shading algorithm matching gradients instead of intensities provides more accurate reconstructions when illumination is incorrectly estimated under a flatness prior.

cs.CV↗

arXiv2018

Critical Contours: An Invariant Linking Image Flow with Salient Surface Organization

We exploit a key result from visual psychophysics---that individuals perceive shape qualitatively---to develop the use of a geometrical/topological "invariant'' (the Morse--Smale complex) relating image structure with surface structure. Differences across individuals are minimal near certain configurations such as ridges and boundaries, and it is these configurations that are often represented in line drawings. In particular, we introduce a method for inferring a qualitative three-dimensional shape from shading patterns that link the shape-from-shading inference with shape-from-contour inference. For a given shape, certain shading patches approach "line drawings'' in a well-defined limit. Under this limit, and invariably with respect to rendering choices, these shading patterns provide a qualitative description of the surface. We further show that, under this model, the contours partition the surface into meaningful parts using the Morse--Smale complex. These critical contours are the (perceptually) stable parts of this complex and are invariant over a wide class of rendering models. Intuitively, our main result shows that critical contours partition smooth surfaces into bumps and valleys, in effect providing a scaffold on the image from which a full surface can be interpolated.

cs.CV↗

arXiv2017

Feature Selection Facilitates Learning Mixtures of Discrete Product Distributions

Feature selection can facilitate the learning of mixtures of discrete random variables as they arise, e.g. in crowdsourcing tasks. Intuitively, not all workers are equally reliable but, if the less reliable ones could be eliminated, then learning should be more robust. By analogy with Gaussian mixture models, we seek a low-order statistical approach, and here introduce an algorithm based on the (pairwise) mutual information. This induces an order over workers that is well structured for the `one coin' model. More generally, it is justified by a goodness-of-fit measure and is validated empirically. Improvement in real data sets can be substantial.

stat.ML↗

arXiv2017

What's In A Patch, I: Tensors, Differential Geometry and Statistical Shading Analysis

cs.CV↗

arXiv2017

Feedback Between Motion and Sensation Provides Nonlinear Boost in Run-and-tumble Navigation

Many organisms navigate gradients by alternating straight motions (runs) with random reorientations (tumbles), transiently suppressing tumbles whenever attractant signal increases. This induces a functional coupling between movement and sensation, since tumbling probability is controlled by the internal state of the organism which, in turn, depends on previous signal levels. Although a negative feedback tends to maintain this internal state close to adapted levels, positive feedback can arise when motion up the gradient reduces tumbling probability, further boosting drift up the gradient. Importantly, such positive feedback can drive large fluctuations in the internal state, complicating analytical approaches. Previous studies focused on what happens when the negative feedback dominates the dynamics. By contrast, we show here that there is a large portion of physiologically-relevant parameter space where the positive feedback can dominate, even when gradients are relatively shallow. We demonstrate how large transients emerge because of non-normal dynamics (non-orthogonal eigenvectors near a stable fixed point) inherent in the positive feedback, and further identify a fundamental nonlinearity that strongly amplifies their effect. Most importantly, this amplification is asymmetric, elongating runs in favorable directions and abbreviating others. The result is a "ratchet-like" gradient climbing behavior with drift speeds that can approach half the maximum run speed of the organism. Our results thus show that the classical drawback of run-and-tumble navigation --- wasteful runs in the wrong direction --- can be mitigated by exploiting the non-normal dynamics implicit in the run-and-tumble strategy.

q-bio.CB↗