arXiv subjects

Andreas S. Tolias

Publications and source records attributed to Andreas S. Tolias.

At least 19 recordsLinked to original sources

arXiv2026

OmniMouse: Scaling properties of multi-modal, multi-task Brain Models on 150B Neural Tokens

Scaling data and artificial neural networks has transformed AI, driving breakthroughs in language and vision. Whether similar principles apply to modeling brain activity remains unclear. Here we leveraged a dataset of 3.1 million neurons from the visual cortex of 73 mice across 323 sessions, totaling more than 150 billion neural tokens recorded during natural movies, images and parametric stimuli, and behavior. We train multi-modal, multi-task models that support three regimes flexibly at test time: neural prediction, behavioral decoding, neural forecasting, or any combination of the three. OmniMouse achieves state-of-the-art performance, outperforming specialized baselines across nearly all evaluation regimes. We find that performance scales reliably with more data, but gains from increasing model size saturate. This inverts the standard AI scaling story: in language and computer vision, massive datasets make parameter scaling the primary driver of progress, whereas in brain modeling -- even in the mouse visual cortex, a relatively simple system -- models remain data-limited despite vast recordings. The observation of systematic scaling raises the possibility of phase transitions in neural modeling, where larger and richer datasets might unlock qualitatively new capabilities, paralleling the emergent properties seen in large language models. Code available at https://github.com/enigma-brain/omnimouse.

q-bio.NC↗

arXiv2026

Letting the neural code speak: Automated characterization of monkey visual neurons through human language

Understanding what individual neurons encode is a core question in neuroscience. In primary visual cortex (V1), mathematical models (e.g., Gabor functions) capture neural selectivity, but no comparable framework exists for higher areas. We show that natural language can fill this role: across macaque V1 and V4, the selectivity of most neurons is captured by concise, verifiable semantic descriptions. Using digital twins of V1 and V4, we develop a closed-loop framework that translates each neuron's high- and low-activating images into dense captions, generates a semantic hypothesis and synthesized images, and verifies the hypothesis in silico. Descriptions range from oriented edges and spatial frequency in V1 to conjunctions of form, color, and texture in V4. In V4, images generated from activating and suppressing hypotheses drove 96.1% of neurons above the 95th and 97.6% below the 5th percentile of natural-image responses, respectively (vs. ~10% for random images); V1 activation results matched V4, while V1 suppression was less describable in language. Representational similarity analysis reveals partial alignment between neural activity, vision embeddings, and language embeddings, with vision most aligned to neural activity; alignment lost in the text bottleneck is recovered when hypotheses are rendered back into images, showing that linguistic compression is lossy yet semantically faithful. Together, these results show that combining generative models with neural digital twins enables interpretable, testable descriptions of neural function at scale, toward agentic scientific discovery.

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arXiv2025

Replay Can Provably Increase Forgetting

Continual learning seeks to enable machine learning systems to solve an increasing corpus of tasks sequentially. A critical challenge for continual learning is forgetting, where the performance on previously learned tasks decreases as new tasks are introduced. One of the commonly used techniques to mitigate forgetting, sample replay, has been shown empirically to reduce forgetting by retaining some examples from old tasks and including them in new training episodes. In this work, we provide a theoretical analysis of sample replay in an over-parameterized continual linear regression setting, where each task is given by a linear subspace and with enough replay samples, one would be able to eliminate forgetting. Our analysis focuses on sample replay and highlights the role of the replayed samples and the relationship between task subspaces. Surprisingly, we find that, even in a noiseless setting, forgetting can be non-monotonic with respect to the number of replay samples. We present tasks where replay can be harmful with respect to worst-case settings, and also in distributional settings where replay of randomly selected samples increases forgetting in expectation. We also give empirical evidence that harmful replay is not limited to training with linear models by showing similar behavior for a neural networks equipped with SGD. Through experiments on a commonly used benchmark, we provide additional evidence that, even in seemingly benign scenarios, performance of the replay heavily depends on the choice of replay samples and the relationship between tasks.

cs.LG↗

arXiv2025

Learning to cluster neuronal function

Deep neural networks trained to predict neural activity from visual input and behaviour have shown great potential to serve as digital twins of the visual cortex. Per-neuron embeddings derived from these models could potentially be used to map the functional landscape or identify cell types. However, state-of-the-art predictive models of mouse V1 do not generate functional embeddings that exhibit clear clustering patterns which would correspond to cell types. This raises the question whether the lack of clustered structure is due to limitations of current models or a true feature of the functional organization of mouse V1. In this work, we introduce DECEMber -- Deep Embedding Clustering via Expectation Maximization-based refinement -- an explicit inductive bias into predictive models that enhances clustering by adding an auxiliary $t$-distribution-inspired loss function that enforces structured organization among per-neuron embeddings. We jointly optimize both neuronal feature embeddings and clustering parameters, updating cluster centers and scale matrices using the EM-algorithm. We demonstrate that these modifications improve cluster consistency while preserving high predictive performance and surpassing standard clustering methods in terms of stability. Moreover, DECEMber generalizes well across species (mice, primates) and visual areas (retina, V1, V4). The code is available at https://github.com/Nisone2000/sensorium/tree/neuroips_version.

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arXiv2024

The Dynamic Sensorium competition for predicting large-scale mouse visual cortex activity from videos

Understanding how biological visual systems process information is challenging due to the complex nonlinear relationship between neuronal responses and high-dimensional visual input. Artificial neural networks have already improved our understanding of this system by allowing computational neuroscientists to create predictive models and bridge biological and machine vision. During the Sensorium 2022, we introduced benchmarks for vision models with static input. However, animals operate and excel in dynamic environments, making it crucial to study and understand how the brain functions under these conditions. Moreover, many biological theories, such as predictive coding, suggest that previous input is crucial for current input processing. Currently, there is no standardized benchmark to identify state-of-the-art dynamic models of the mouse visual system. To address this gap, we propose the Sensorium 2023 Benchmark Competition with dynamic input. It includes the collection of a new large-scale dataset from the primary visual cortex of ten mice, containing responses from over 78,000 neurons to over 2 hours of dynamic stimuli per neuron. Participants in the main benchmark track will compete to identify the best predictive models of neuronal responses for dynamic input. We will also host a bonus track in which submission performance will be evaluated on out-of-domain input, using withheld neuronal responses to dynamic input stimuli whose statistics differ from the training set. Both tracks will offer behavioral data along with video stimuli. As before, we will provide code, tutorials, and strong pre-trained baseline models to encourage participation. We hope this competition will continue to strengthen the accompanying Sensorium benchmarks collection as a standard tool to measure progress in large-scale neural system identification models of the entire mouse visual hierarchy and beyond.

q-bio.NC↗

arXiv2024

Retrospective for the Dynamic Sensorium Competition for predicting large-scale mouse primary visual cortex activity from videos

Understanding how biological visual systems process information is challenging because of the nonlinear relationship between visual input and neuronal responses. Artificial neural networks allow computational neuroscientists to create predictive models that connect biological and machine vision. Machine learning has benefited tremendously from benchmarks that compare different model on the same task under standardized conditions. However, there was no standardized benchmark to identify state-of-the-art dynamic models of the mouse visual system. To address this gap, we established the Sensorium 2023 Benchmark Competition with dynamic input, featuring a new large-scale dataset from the primary visual cortex of ten mice. This dataset includes responses from 78,853 neurons to 2 hours of dynamic stimuli per neuron, together with the behavioral measurements such as running speed, pupil dilation, and eye movements. The competition ranked models in two tracks based on predictive performance for neuronal responses on a held-out test set: one focusing on predicting in-domain natural stimuli and another on out-of-distribution (OOD) stimuli to assess model generalization. As part of the NeurIPS 2023 competition track, we received more than 160 model submissions from 22 teams. Several new architectures for predictive models were proposed, and the winning teams improved the previous state-of-the-art model by 50%. Access to the dataset as well as the benchmarking infrastructure will remain online at www.sensorium-competition.net.

q-bio.NC↗

arXiv2024

Continual Learning: Applications and the Road Forward

Continual learning is a subfield of machine learning, which aims to allow machine learning models to continuously learn on new data, by accumulating knowledge without forgetting what was learned in the past. In this work, we take a step back, and ask: "Why should one care about continual learning in the first place?". We set the stage by examining recent continual learning papers published at four major machine learning conferences, and show that memory-constrained settings dominate the field. Then, we discuss five open problems in machine learning, and even though they might seem unrelated to continual learning at first sight, we show that continual learning will inevitably be part of their solution. These problems are model editing, personalization and specialization, on-device learning, faster (re-)training and reinforcement learning. Finally, by comparing the desiderata from these unsolved problems and the current assumptions in continual learning, we highlight and discuss four future directions for continual learning research. We hope that this work offers an interesting perspective on the future of continual learning, while displaying its potential value and the paths we have to pursue in order to make it successful. This work is the result of the many discussions the authors had at the Dagstuhl seminar on Deep Continual Learning, in March 2023.

cs.LG↗

arXiv2024

Most discriminative stimuli for functional cell type clustering

Identifying cell types and understanding their functional properties is crucial for unraveling the mechanisms underlying perception and cognition. In the retina, functional types can be identified by carefully selected stimuli, but this requires expert domain knowledge and biases the procedure towards previously known cell types. In the visual cortex, it is still unknown what functional types exist and how to identify them. Thus, for unbiased identification of the functional cell types in retina and visual cortex, new approaches are needed. Here we propose an optimization-based clustering approach using deep predictive models to obtain functional clusters of neurons using Most Discriminative Stimuli (MDS). Our approach alternates between stimulus optimization with cluster reassignment akin to an expectation-maximization algorithm. The algorithm recovers functional clusters in mouse retina, marmoset retina and macaque visual area V4. This demonstrates that our approach can successfully find discriminative stimuli across species, stages of the visual system and recording techniques. The resulting most discriminative stimuli can be used to assign functional cell types fast and on the fly, without the need to train complex predictive models or show a large natural scene dataset, paving the way for experiments that were previously limited by experimental time. Crucially, MDS are interpretable: they visualize the distinctive stimulus patterns that most unambiguously identify a specific type of neuron.

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arXiv2023

Generalization properties of contrastive world models

Recent work on object-centric world models aim to factorize representations in terms of objects in a completely unsupervised or self-supervised manner. Such world models are hypothesized to be a key component to address the generalization problem. While self-supervision has shown improved performance however, OOD generalization has not been systematically and explicitly tested. In this paper, we conduct an extensive study on the generalization properties of contrastive world model. We systematically test the model under a number of different OOD generalization scenarios such as extrapolation to new object attributes, introducing new conjunctions or new attributes. Our experiments show that the contrastive world model fails to generalize under the different OOD tests and the drop in performance depends on the extent to which the samples are OOD. When visualizing the transition updates and convolutional feature maps, we observe that any changes in object attributes (such as previously unseen colors, shapes, or conjunctions of color and shape) breaks down the factorization of object representations. Overall, our work highlights the importance of object-centric representations for generalization and current models are limited in their capacity to learn such representations required for human-level generalization.

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arXiv2023

Toward Next-Generation Artificial Intelligence: Catalyzing the NeuroAI Revolution

Neuroscience has long been an essential driver of progress in artificial intelligence (AI). We propose that to accelerate progress in AI, we must invest in fundamental research in NeuroAI. A core component of this is the embodied Turing test, which challenges AI animal models to interact with the sensorimotor world at skill levels akin to their living counterparts. The embodied Turing test shifts the focus from those capabilities like game playing and language that are especially well-developed or uniquely human to those capabilities, inherited from over 500 million years of evolution, that are shared with all animals. Building models that can pass the embodied Turing test will provide a roadmap for the next generation of AI.

cs.AI↗

arXiv2022

Learning Dynamics and Structure of Complex Systems Using Graph Neural Networks

Many complex systems are composed of interacting parts, and the underlying laws are usually simple and universal. While graph neural networks provide a useful relational inductive bias for modeling such systems, generalization to new system instances of the same type is less studied. In this work we trained graph neural networks to fit time series from an example nonlinear dynamical system, the belief propagation algorithm. We found simple interpretations of the learned representation and model components, and they are consistent with core properties of the probabilistic inference algorithm. We successfully identified a 'graph translator' between the statistical interactions in belief propagation and parameters of the corresponding trained network, and showed that it enables two types of novel generalization: to recover the underlying structure of a new system instance based solely on time series observations, or to construct a new network from this structure directly. Our results demonstrated a path towards understanding both dynamics and structure of a complex system and how such understanding can be used for generalization.

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arXiv2022

The Sensorium competition on predicting large-scale mouse primary visual cortex activity

The neural underpinning of the biological visual system is challenging to study experimentally, in particular as the neuronal activity becomes increasingly nonlinear with respect to visual input. Artificial neural networks (ANNs) can serve a variety of goals for improving our understanding of this complex system, not only serving as predictive digital twins of sensory cortex for novel hypothesis generation in silico, but also incorporating bio-inspired architectural motifs to progressively bridge the gap between biological and machine vision. The mouse has recently emerged as a popular model system to study visual information processing, but no standardized large-scale benchmark to identify state-of-the-art models of the mouse visual system has been established. To fill this gap, we propose the Sensorium benchmark competition. We collected a large-scale dataset from mouse primary visual cortex containing the responses of more than 28,000 neurons across seven mice stimulated with thousands of natural images, together with simultaneous behavioral measurements that include running speed, pupil dilation, and eye movements. The benchmark challenge will rank models based on predictive performance for neuronal responses on a held-out test set, and includes two tracks for model input limited to either stimulus only (Sensorium) or stimulus plus behavior (Sensorium+). We provide a starting kit to lower the barrier for entry, including tutorials, pre-trained baseline models, and APIs with one line commands for data loading and submission. We would like to see this as a starting point for regular challenges and data releases, and as a standard tool for measuring progress in large-scale neural system identification models of the mouse visual system and beyond.

q-bio.NC↗

arXiv2022

Probing Few-Shot Generalization with Attributes

Despite impressive progress in deep learning, generalizing far beyond the training distribution is an important open challenge. In this work, we consider few-shot classification, and aim to shed light on what makes some novel classes easier to learn than others, and what types of learned representations generalize better. To this end, we define a new paradigm in terms of attributes -- simple building blocks of which concepts are formed -- as a means of quantifying the degree of relatedness of different concepts. Our empirical analysis reveals that supervised learning generalizes poorly to new attributes, but a combination of self-supervised pretraining with supervised finetuning leads to stronger generalization. The benefit of self-supervised pretraining and supervised finetuning is further investigated through controlled experiments using random splits of the attribute space, and we find that predictability of test attributes provides an informative estimate of a model's generalization ability.

cs.LG↗

arXiv2022

Understanding robustness and generalization of artificial neural networks through Fourier masks

Despite the enormous success of artificial neural networks (ANNs) in many disciplines, the characterization of their computations and the origin of key properties such as generalization and robustness remain open questions. Recent literature suggests that robust networks with good generalization properties tend to be biased towards processing low frequencies in images. To explore the frequency bias hypothesis further, we develop an algorithm that allows us to learn modulatory masks highlighting the essential input frequencies needed for preserving a trained network's performance. We achieve this by imposing invariance in the loss with respect to such modulations in the input frequencies. We first use our method to test the low-frequency preference hypothesis of adversarially trained or data-augmented networks. Our results suggest that adversarially robust networks indeed exhibit a low-frequency bias but we find this bias is also dependent on directions in frequency space. However, this is not necessarily true for other types of data augmentation. Our results also indicate that the essential frequencies in question are effectively the ones used to achieve generalization in the first place. Surprisingly, images seen through these modulatory masks are not recognizable and resemble texture-like patterns.

cs.CV↗

arXiv2021

Towards robust vision by multi-task learning on monkey visual cortex

Deep neural networks set the state-of-the-art across many tasks in computer vision, but their generalization ability to image distortions is surprisingly fragile. In contrast, the mammalian visual system is robust to a wide range of perturbations. Recent work suggests that this generalization ability can be explained by useful inductive biases encoded in the representations of visual stimuli throughout the visual cortex. Here, we successfully leveraged these inductive biases with a multi-task learning approach: we jointly trained a deep network to perform image classification and to predict neural activity in macaque primary visual cortex (V1). We measured the out-of-distribution generalization abilities of our network by testing its robustness to image distortions. We found that co-training on monkey V1 data leads to increased robustness despite the absence of those distortions during training. Additionally, we showed that our network's robustness is very close to that of an Oracle network where parts of the architecture are directly trained on noisy images. Our results also demonstrated that the network's representations become more brain-like as their robustness improves. Using a novel constrained reconstruction analysis, we investigated what makes our brain-regularized network more robust. We found that our co-trained network is more sensitive to content than noise when compared to a Baseline network that we trained for image classification alone. Using DeepGaze-predicted saliency maps for ImageNet images, we found that our monkey co-trained network tends to be more sensitive to salient regions in a scene, reminiscent of existing theories on the role of V1 in the detection of object borders and bottom-up saliency. Overall, our work expands the promising research avenue of transferring inductive biases from the brain, and provides a novel analysis of the effects of our transfer.

cs.CV↗

arXiv2021

Class-Incremental Learning with Generative Classifiers

Incrementally training deep neural networks to recognize new classes is a challenging problem. Most existing class-incremental learning methods store data or use generative replay, both of which have drawbacks, while 'rehearsal-free' alternatives such as parameter regularization or bias-correction methods do not consistently achieve high performance. Here, we put forward a new strategy for class-incremental learning: generative classification. Rather than directly learning the conditional distribution p(y|x), our proposal is to learn the joint distribution p(x,y), factorized as p(x|y)p(y), and to perform classification using Bayes' rule. As a proof-of-principle, here we implement this strategy by training a variational autoencoder for each class to be learned and by using importance sampling to estimate the likelihoods p(x|y). This simple approach performs very well on a diverse set of continual learning benchmarks, outperforming generative replay and other existing baselines that do not store data.

cs.LG↗

arXiv2020

Factorized Neural Processes for Neural Processes: $K$-Shot Prediction of Neural Responses

In recent years, artificial neural networks have achieved state-of-the-art performance for predicting the responses of neurons in the visual cortex to natural stimuli. However, they require a time consuming parameter optimization process for accurately modeling the tuning function of newly observed neurons, which prohibits many applications including real-time, closed-loop experiments. We overcome this limitation by formulating the problem as $K$-shot prediction to directly infer a neuron's tuning function from a small set of stimulus-response pairs using a Neural Process. This required us to developed a Factorized Neural Process, which embeds the observed set into a latent space partitioned into the receptive field location and the tuning function properties. We show on simulated responses that the predictions and reconstructed receptive fields from the Factorized Neural Process approach ground truth with increasing number of trials. Critically, the latent representation that summarizes the tuning function of a neuron is inferred in a quick, single forward pass through the network. Finally, we validate this approach on real neural data from visual cortex and find that the predictive accuracy is comparable to -- and for small $K$ even greater than -- optimization based approaches, while being substantially faster. We believe this novel deep learning systems identification framework will facilitate better real-time integration of artificial neural network modeling into neuroscience experiments.

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arXiv2019

Learning From Brains How to Regularize Machines

Despite impressive performance on numerous visual tasks, Convolutional Neural Networks (CNNs) --- unlike brains --- are often highly sensitive to small perturbations of their input, e.g. adversarial noise leading to erroneous decisions. We propose to regularize CNNs using large-scale neuroscience data to learn more robust neural features in terms of representational similarity. We presented natural images to mice and measured the responses of thousands of neurons from cortical visual areas. Next, we denoised the notoriously variable neural activity using strong predictive models trained on this large corpus of responses from the mouse visual system, and calculated the representational similarity for millions of pairs of images from the model's predictions. We then used the neural representation similarity to regularize CNNs trained on image classification by penalizing intermediate representations that deviated from neural ones. This preserved performance of baseline models when classifying images under standard benchmarks, while maintaining substantially higher performance compared to baseline or control models when classifying noisy images. Moreover, the models regularized with cortical representations also improved model robustness in terms of adversarial attacks. This demonstrates that regularizing with neural data can be an effective tool to create an inductive bias towards more robust inference.

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