Jean-Rémi King

Yohann Benchetrit, Marlène Careil, Simon Dahan et al.

Brain decoding is limited by the availability of labeled neural data, and remains challenging in low-data regimes. To address this issue, we investigate whether and when brain decoding can be boosted by augmenting small fMRI datasets with synthetic data generated by a pretrained model of fMRI responses to stimuli. We use TRIBE v2, a large encoding model pretrained on more than 1000 hours of fMRI responses to video, audio and language. For each dataset, we evaluate systematic grids that show how the performance of image decoders varies with the amount of synthetic data used for training. Our results, based on two datasets (the 7T fMRI Natural Scenes Dataset and 3T fMRI BOLD5000), show up to 68% improvement in Top-10 image-retrieval accuracy compared to decoders trained only on real data. Importantly, the proportion of augmented data required to reach a given image decoding performance needs to be adjusted depending on the data source. Surprisingly, image decoders trained exclusively on synthetic fMRI can perform above chance in some settings, suggesting that TRIBE v2 can support zero-shot brain-to-image decoding. Together, these results show how large-scale models of the fMRI responses to sight, sound and language may provide a foundation to improve the data efficiency for image decoding.

Alexandre Défossez, Charlotte Caucheteux, Jérémy Rapin et al.

Decoding speech from brain activity is a long-awaited goal in both healthcare and neuroscience. Invasive devices have recently led to major milestones in that regard: deep learning algorithms trained on intracranial recordings now start to decode elementary linguistic features (e.g. letters, words, spectrograms). However, extending this approach to natural speech and non-invasive brain recordings remains a major challenge. Here, we introduce a model trained with contrastive-learning to decode self-supervised representations of perceived speech from the non-invasive recordings of a large cohort of healthy individuals. To evaluate this approach, we curate and integrate four public datasets, encompassing 175 volunteers recorded with magneto- or electro-encephalography (M/EEG), while they listened to short stories and isolated sentences. The results show that our model can identify, from 3 seconds of MEG signals, the corresponding speech segment with up to 41% accuracy out of more than 1,000 distinct possibilities on average across participants, and more than 80% in the very best participants - a performance that allows the decoding of words and phrases absent from the training set. The comparison of our model to a variety of baselines highlights the importance of (i) a contrastive objective, (ii) pretrained representations of speech and (iii) a common convolutional architecture simultaneously trained across multiple participants. Finally, the analysis of the decoder's predictions suggests that they primarily depend on lexical and contextual semantic representations. Overall, this effective decoding of perceived speech from non-invasive recordings delineates a promising path to decode language from brain activity, without putting patients at risk for brain surgery.

Linnea Evanson, Yair Lakretz, Jean-Rémi King

During language acquisition, children follow a typical sequence of learning stages, whereby they first learn to categorize phonemes before they develop their lexicon and eventually master increasingly complex syntactic structures. However, the computational principles that lead to this learning trajectory remain largely unknown. To investigate this, we here compare the learning trajectories of deep language models to those of children. Specifically, we test whether, during its training, GPT-2 exhibits stages of language acquisition comparable to those observed in children aged between 18 months and 6 years. For this, we train 48 GPT-2 models from scratch and evaluate their syntactic and semantic abilities at each training step, using 96 probes curated from the BLiMP, Zorro and BIG-Bench benchmarks. We then compare these evaluations with the behavior of 54 children during language production. Our analyses reveal three main findings. First, similarly to children, the language models tend to learn linguistic skills in a systematic order. Second, this learning scheme is parallel: the language tasks that are learned last improve from the very first training steps. Third, some - but not all - learning stages are shared between children and these language models. Overall, these results shed new light on the principles of language acquisition, and highlight important divergences in how humans and modern algorithms learn to process natural language.

Stéphane d'Ascoli, Jérémy Rapin, Yohann Benchetrit et al.

Cognitive neuroscience is fragmented into specialized models, each tailored to specific experimental paradigms, hence preventing a unified model of cognition in the human brain. Here, we introduce TRIBE v2, a tri-modal (video, audio and language) foundation model capable of predicting human brain activity in a variety of naturalistic and experimental conditions. Leveraging a unified dataset of over 1,000 hours of fMRI across 720 subjects, we demonstrate that our model accurately predicts high-resolution brain responses for novel stimuli, tasks and subjects, superseding traditional linear encoding models, delivering several-fold improvements in accuracy. Critically, TRIBE v2 enables in silico experimentation: tested on seminal visual and neuro-linguistic paradigms, it recovers a variety of results established by decades of empirical research. Finally, by extracting interpretable latent features, TRIBE v2 reveals the fine-grained topography of multisensory integration. These results establish artificial intelligence as a unifying framework for exploring the functional organization of the human brain.

Yohann Benchetrit, Hubert Banville, Jean-Rémi King

In the past five years, the use of generative and foundational AI systems has greatly improved the decoding of brain activity. Visual perception, in particular, can now be decoded from functional Magnetic Resonance Imaging (fMRI) with remarkable fidelity. This neuroimaging technique, however, suffers from a limited temporal resolution ($\approx$0.5 Hz) and thus fundamentally constrains its real-time usage. Here, we propose an alternative approach based on magnetoencephalography (MEG), a neuroimaging device capable of measuring brain activity with high temporal resolution ($\approx$5,000 Hz). For this, we develop an MEG decoding model trained with both contrastive and regression objectives and consisting of three modules: i) pretrained embeddings obtained from the image, ii) an MEG module trained end-to-end and iii) a pretrained image generator. Our results are threefold: Firstly, our MEG decoder shows a 7X improvement of image-retrieval over classic linear decoders. Second, late brain responses to images are best decoded with DINOv2, a recent foundational image model. Third, image retrievals and generations both suggest that high-level visual features can be decoded from MEG signals, although the same approach applied to 7T fMRI also recovers better low-level features. Overall, these results, while preliminary, provide an important step towards the decoding -- in real-time -- of the visual processes continuously unfolding within the human brain.

Joséphine Raugel, Maximilian Seitzer, Marc Szafraniec et al.

Backpropagation is the core learning mechanism underlying deep learning. However, whether and how this algorithm is implemented in the brain remains highly debated. In particular, while forward activations of pretrained models reliably map onto the cortical hierarchy of visual processing, it is unknown whether backpropagated gradients exhibit a similar correspondence. Here, we address this question using functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG) recordings of human brain responses to natural images. For this, we extend standard encoding analyses of forward activations to map backpropagated gradients onto neural data. Focusing on a recent self-supervised vision model (DINOv3) and reproducing results on eight vision models, we find that backpropagated gradients can reliably predict both fMRI and MEG signals, specifically in higher-level visual cortex and for later latencies. However, the spatial and temporal organization of these backpropagated gradients in the brain diverges from the patterns expected under a biologically plausible backpropagation mechanism: specifically, both the order in which gradients are computed and their spatial organization diverge from the temporal and spatial hierarchies of the human brain. Together, these results suggest that, although deep networks and the brain may share similar representational content, they likely rely on fundamentally different mechanisms to learn those representations.

Hubert Banville, Stéphane d'Ascoli, Simon Dahan et al.

Deep learning and large public datasets have recently catalyzed the proliferation of AI models for processing brain recordings. However, systematically evaluating these models remains a challenge: not only do the preprocessing pipelines, training and finetuning approaches largely vary across studies, but their downstream evaluation is often limited to small sets of tasks and/or datasets. Here, we present NeuralBench: a unified framework for benchmarking AI models of brain activity. We accompany this framework with NeuralBench-EEG v1.0 -- a large EEG benchmark that includes 36 electroencephalography (EEG) tasks and 14 deep learning architectures, and is evaluated on 94 datasets accessed through a standardized interface. This first EEG-focused release already highlights two main findings. First, current foundation models only marginally outperform task-specific models. Second, a large set of tasks (e.g. cognitive decoding, clinical predictions) remain highly challenging, even for the best models. Critically, NeuralBench is designed for the integration of new tasks, datasets, models, and neuroimaging modalities, as illustrated by preliminary extensions to MEG and fMRI datasets and models. Through this white paper, we invite the community to expand this open-source framework and work together toward a unified benchmarking standard for neuroimaging models.

Pierre Orhan, Pablo Diego-Simón, Emmnanuel Chemla et al.

During language acquisition, children successively learn to categorize phonemes, identify words, and combine them with syntax to form new meaning. While the development of this behavior is well characterized, we still lack a unifying computational framework to explain its underlying neural representations. Here, we investigate whether and when phonemic, lexical, and syntactic representations emerge in the activations of artificial neural networks during their training. Our results show that both speech- and text-based models follow a sequence of learning stages: during training, their neural activations successively build subspaces, where the geometry of the neural activations represents phonemic, lexical, and syntactic structure. While this developmental trajectory qualitatively relates to children's, it is quantitatively different: These algorithms indeed require two to four orders of magnitude more data for these neural representations to emerge. Together, these results show conditions under which major stages of language acquisition spontaneously emerge, and hence delineate a promising path to understand the computations underpinning language acquisition.

Linnea Evanson, Mingfang, Zhang et al.

Decoding speech from brain activity has typically relied on limited neural recordings collected during short and highly controlled experiments. Here, we introduce a framework to leverage week-long intracranial and audio recordings from patients undergoing clinical monitoring, effectively increasing the training dataset size by over two orders of magnitude. With this pretraining, our contrastive learning model substantially outperforms models trained solely on classic experimental data, with gains that scale log-linearly with dataset size. Analysis of the learned representations reveals that, while brain activity represents speech features, its global structure largely drifts across days, highlighting the need for models that explicitly account for cross-day variability. Overall, our approach opens a scalable path toward decoding and modeling brain representations in both real-life and controlled task settings.

Stéphane d'Ascoli, Jérémy Rapin, Yohann Benchetrit et al.

Historically, neuroscience has progressed by fragmenting into specialized domains, each focusing on isolated modalities, tasks, or brain regions. While fruitful, this approach hinders the development of a unified model of cognition. Here, we introduce TRIBE, the first deep neural network trained to predict brain responses to stimuli across multiple modalities, cortical areas and individuals. By combining the pretrained representations of text, audio and video foundational models and handling their time-evolving nature with a transformer, our model can precisely model the spatial and temporal fMRI responses to videos, achieving the first place in the Algonauts 2025 brain encoding competition with a significant margin over competitors. Ablations show that while unimodal models can reliably predict their corresponding cortical networks (e.g. visual or auditory networks), they are systematically outperformed by our multimodal model in high-level associative cortices. Currently applied to perception and comprehension, our approach paves the way towards building an integrative model of representations in the human brain. Our code is available at https://github.com/facebookresearch/algonauts-2025.

Pablo J. Diego-Simón, Pierre Orhan, Yair Lakretz et al.

How do artificial neural networks bind concepts to form complex semantic structures? Here, we propose a simple neural code, whereby the existence and the type of relations between entities are represented by the distance and the direction between their embeddings, respectively. We test this hypothesis in a variety of Large Language Models (LLMs), each input with natural-language descriptions of minimalist tasks from five different domains: arithmetic, visual scenes, family trees, metro maps and social interactions. Results show that the true semantic structures can be linearly recovered with a Polar Probe targeting a subspace of LLMs' layer activations. Second, this code emerges mostly in middle layers and improves with LLM performance. Third, these Polar Probes successfully generalize to new entities and relation types, but degrades with the size of the semantic structure. Finally, the quality of the polar representation correlates with the LLM's ability to answer questions about the semantic structure. Together, these findings suggest that LLMs learn to build complex semantic structures by binding representations with a simple geometrical principle.

Joséphine Raugel, Stéphane d'Ascoli, Jérémy Rapin et al.

Recent studies suggest that the representations learned by large language models (LLMs) are partially aligned to those of the human brain. However, whether and why this alignment score arises from a similar sequence of computations remains elusive. In this study, we explore this question by examining temporally-resolved brain signals of participants listening to 10 hours of an audiobook. We study these neural dynamics jointly with a benchmark encompassing 22 LLMs varying in size and architecture type. Our analyses confirm that LLMs and the brain generate representations in a similar order: specifically, activations in the initial layers of LLMs tend to best align with early brain responses, while the deeper layers of LLMs tend to best align with later brain responses. This brain-LLM alignment is consistent across transformers and recurrent architectures. However, its emergence depends on both model size and context length. Overall, this study sheds light on the sequential nature of computations and the factors underlying the partial convergence between biological and artificial neural networks.

Jarod Lévy, Mingfang Zhang, Svetlana Pinet et al.

Modern neuroprostheses can now restore communication in patients who have lost the ability to speak or move. However, these invasive devices entail risks inherent to neurosurgery. Here, we introduce a non-invasive method to decode the production of sentences from brain activity and demonstrate its efficacy in a cohort of 35 healthy volunteers. For this, we present Brain2Qwerty, a new deep learning architecture trained to decode sentences from either electro- (EEG) or magneto-encephalography (MEG), while participants typed briefly memorized sentences on a QWERTY keyboard. With MEG, Brain2Qwerty reaches, on average, a character-error-rate (CER) of 32% and substantially outperforms EEG (CER: 67%). For the best participants, the model achieves a CER of 19%, and can perfectly decode a variety of sentences outside of the training set. While error analyses suggest that decoding depends on motor processes, the analysis of typographical errors suggests that it also involves higher-level cognitive factors. Overall, these results narrow the gap between invasive and non-invasive methods and thus open the path for developing safe brain-computer interfaces for non-communicating patients.

Stéphane d'Ascoli, Corentin Bel, Jérémy Rapin et al.

Deep learning has recently enabled the decoding of language from the neural activity of a few participants with electrodes implanted inside their brain. However, reliably decoding words from non-invasive recordings remains an open challenge. To tackle this issue, we introduce a novel deep learning pipeline to decode individual words from non-invasive electro- (EEG) and magneto-encephalography (MEG) signals. We train and evaluate our approach on an unprecedentedly large number of participants (723) exposed to five million words either written or spoken in English, French or Dutch. Our model outperforms existing methods consistently across participants, devices, languages, and tasks, and can decode words absent from the training set. Our analyses highlight the importance of the recording device and experimental protocol: MEG and reading are easier to decode than EEG and listening, respectively, and it is preferable to collect a large amount of data per participant than to repeat stimuli across a large number of participants. Furthermore, decoding performance consistently increases with the amount of (i) data used for training and (ii) data used for averaging during testing. Finally, single-word predictions show that our model effectively relies on word semantics but also captures syntactic and surface properties such as part-of-speech, word length and even individual letters, especially in the reading condition. Overall, our findings delineate the path and remaining challenges towards building non-invasive brain decoders for natural language.

Alexis Thual, Yohann Benchetrit, Felix Geilert et al.

Deep learning is leading to major advances in the realm of brain decoding from functional Magnetic Resonance Imaging (fMRI). However, the large inter-subject variability in brain characteristics has limited most studies to train models on one subject at a time. Consequently, this approach hampers the training of deep learning models, which typically requires very large datasets. Here, we propose to boost brain decoding by aligning brain responses to videos and static images across subjects. Compared to the anatomically-aligned baseline, our method improves out-of-subject decoding performance by up to 75%. Moreover, it also outperforms classical single-subject approaches when fewer than 100 minutes of data is available for the tested subject. Furthermore, we propose a new multi-subject alignment method, which obtains comparable results to that of classical single-subject approaches while improving out-of-subject generalization. Finally, we show that this method aligns neural representations in accordance with brain anatomy. Overall, this study lays the foundations for leveraging extensive neuroimaging datasets and enhancing the decoding of individuals with a limited amount of brain recordings.

Hubert Banville, Yohann Benchetrit, Stéphane d'Ascoli et al.

Generative AI has recently propelled the decoding of images from brain activity. How do these approaches scale with the amount and type of neural recordings? Here, we systematically compare image decoding from four types of non-invasive devices: electroencephalography (EEG), magnetoencephalography (MEG), high-field functional Magnetic Resonance Imaging (3T fMRI) and ultra-high field (7T) fMRI. For this, we evaluate decoding models on the largest benchmark to date, encompassing 8 public datasets, 84 volunteers, 498 hours of brain recording and 2.3 million brain responses to natural images. Unlike previous work, we focus on single-trial decoding performance to simulate real-time settings. This systematic comparison reveals three main findings. First, the most precise neuroimaging devices tend to yield the best decoding performances, when the size of the training sets are similar. However, the gain enabled by deep learning - in comparison to linear models - is obtained with the noisiest devices. Second, we do not observe any plateau of decoding performance as the amount of training data increases. Rather, decoding performance scales log-linearly with the amount of brain recording. Third, this scaling law primarily depends on the amount of data per subject. However, little decoding gain is observed by increasing the number of subjects. Overall, these findings delineate the path most suitable to scale the decoding of images from non-invasive brain recordings.

Joséphine Raugel, Marc Szafraniec, Huy V. Vo et al.

Many AI models trained on natural images develop representations that resemble those of the human brain. However, the factors that drive this brain-model similarity remain poorly understood. To disentangle how the model, training and data independently lead a neural network to develop brain-like representations, we trained a family of self-supervised vision transformers (DINOv3) that systematically varied these different factors. We compare their representations of images to those of the human brain recorded with both fMRI and MEG, providing high resolution in spatial and temporal analyses. We assess the brain-model similarity with three complementary metrics focusing on overall representational similarity, topographical organization, and temporal dynamics. We show that all three factors - model size, training amount, and image type - independently and interactively impact each of these brain similarity metrics. In particular, the largest DINOv3 models trained with the most human-centric images reach the highest brain-similarity. This emergence of brain-like representations in AI models follows a specific chronology during training: models first align with the early representations of the sensory cortices, and only align with the late and prefrontal representations of the brain with considerably more training. Finally, this developmental trajectory is indexed by both structural and functional properties of the human cortex: the representations that are acquired last by the models specifically align with the cortical areas with the largest developmental expansion, thickness, least myelination, and slowest timescales. Overall, these findings disentangle the interplay between architecture and experience in shaping how artificial neural networks come to see the world as humans do, thus offering a promising framework to understand how the human brain comes to represent its visual world.

Pablo Diego-Simón, Stéphane D'Ascoli, Emmanuel Chemla et al.

Originally formalized with symbolic representations, syntactic trees may also be effectively represented in the activations of large language models (LLMs). Indeed, a 'Structural Probe' can find a subspace of neural activations, where syntactically related words are relatively close to one-another. However, this syntactic code remains incomplete: the distance between the Structural Probe word embeddings can represent the existence but not the type and direction of syntactic relations. Here, we hypothesize that syntactic relations are, in fact, coded by the relative direction between nearby embeddings. To test this hypothesis, we introduce a 'Polar Probe' trained to read syntactic relations from both the distance and the direction between word embeddings. Our approach reveals three main findings. First, our Polar Probe successfully recovers the type and direction of syntactic relations, and substantially outperforms the Structural Probe by nearly two folds. Second, we confirm that this polar coordinate system exists in a low-dimensional subspace of the intermediate layers of many LLMs and becomes increasingly precise in the latest frontier models. Third, we demonstrate with a new benchmark that similar syntactic relations are coded similarly across the nested levels of syntactic trees. Overall, this work shows that LLMs spontaneously learn a geometry of neural activations that explicitly represents the main symbolic structures of linguistic theory.

Pablo J. Diego-Simón, Emmanuel Chemla, Jean-Rémi King et al.

The syntactic structures of sentences can be readily read-out from the activations of large language models (LLMs). However, the ``structural probes'' that have been developed to reveal this phenomenon are typically evaluated on an indiscriminate set of sentences. Consequently, it remains unclear whether structural and/or statistical factors systematically affect these syntactic representations. To address this issue, we conduct an in-depth analysis of structural probes on three controlled benchmarks. Our results are three-fold. First, structural probes are biased by a superficial property: the closer two words are in a sentence, the more likely structural probes will consider them as syntactically linked. Second, structural probes are challenged by linguistic properties: they poorly represent deep syntactic structures, and get interfered by interacting nouns or ungrammatical verb forms. Third, structural probes do not appear to be affected by the predictability of individual words. Overall, this work sheds light on the current challenges faced by structural probes. Providing a benchmark made of controlled stimuli to better evaluate their performance.

Marlène Careil, Yohann Benchetrit, Jean-Rémi King

Brain-to-image decoding has been recently propelled by the progress in generative AI models and the availability of large ultra-high field functional Magnetic Resonance Imaging (fMRI). However, current approaches depend on complicated multi-stage pipelines and preprocessing steps that typically collapse the temporal dimension of brain recordings, thereby limiting time-resolved brain decoders. Here, we introduce Dynadiff (Dynamic Neural Activity Diffusion for Image Reconstruction), a new single-stage diffusion model designed for reconstructing images from dynamically evolving fMRI recordings. Our approach offers three main contributions. First, Dynadiff simplifies training as compared to existing approaches. Second, our model outperforms state-of-the-art models on time-resolved fMRI signals, especially on high-level semantic image reconstruction metrics, while remaining competitive on preprocessed fMRI data that collapse time. Third, this approach allows a precise characterization of the evolution of image representations in brain activity. Overall, this work lays the foundation for time-resolved brain-to-image decoding.

Pierre Orhan, Yves Boubenec, Jean-Rémi King

Over the last decade, numerous studies have shown that deep neural networks exhibit sensory representations similar to those of the mammalian brain, in that their activations linearly map onto cortical responses to the same sensory inputs. However, it remains unknown whether these artificial networks also learn like the brain. To address this issue, we analyze the brain responses of two ferret auditory cortices recorded with functional UltraSound imaging (fUS), while the animals were presented with 320 10\,s sounds. We compare these brain responses to the activations of Wav2vec 2.0, a self-supervised neural network pretrained with 960\,h of speech, and input with the same 320 sounds. Critically, we evaluate Wav2vec 2.0 under two distinct modes: (i) "Pretrained", where the same model is used for all sounds, and (ii) "Continuous Update", where the weights of the pretrained model are modified with back-propagation after every sound, presented in the same order as the ferrets. Our results show that the Continuous-Update mode leads Wav2Vec 2.0 to generate activations that are more similar to the brain than a Pretrained Wav2Vec 2.0 or than other control models using different training modes. These results suggest that the trial-by-trial modifications of self-supervised algorithms induced by back-propagation aligns with the corresponding fluctuations of cortical responses to sounds. Our finding thus provides empirical evidence of a common learning mechanism between self-supervised models and the mammalian cortex during sound processing.

Charlotte Caucheteux, Alexandre Gramfort, Jean-Rémi King

A popular approach to decompose the neural bases of language consists in correlating, across individuals, the brain responses to different stimuli (e.g. regular speech versus scrambled words, sentences, or paragraphs). Although successful, this `model-free' approach necessitates the acquisition of a large and costly set of neuroimaging data. Here, we show that a model-based approach can reach equivalent results within subjects exposed to natural stimuli. We capitalize on the recently-discovered similarities between deep language models and the human brain to compute the mapping between i) the brain responses to regular speech and ii) the activations of deep language models elicited by modified stimuli (e.g. scrambled words, sentences, or paragraphs). Our model-based approach successfully replicates the seminal study of Lerner et al. (2011), which revealed the hierarchy of language areas by comparing the functional-magnetic resonance imaging (fMRI) of seven subjects listening to 7min of both regular and scrambled narratives. We further extend and precise these results to the brain signals of 305 individuals listening to 4.1 hours of narrated stories. Overall, this study paves the way for efficient and flexible analyses of the brain bases of language.

Yair Lakretz, Théo Desbordes, Jean-Rémi King et al.

One of the fundamental principles of contemporary linguistics states that language processing requires the ability to extract recursively nested tree structures. However, it remains unclear whether and how this code could be implemented in neural circuits. Recent advances in Recurrent Neural Networks (RNNs), which achieve near-human performance in some language tasks, provide a compelling model to address such questions. Here, we present a new framework to study recursive processing in RNNs, using subject-verb agreement as a probe into the representations of the neural network. We trained six distinct types of RNNs on a simplified probabilistic context-free grammar designed to independently manipulate the length of a sentence and the depth of its syntactic tree. All RNNs generalized to subject-verb dependencies longer than those seen during training. However, none systematically generalized to deeper tree structures, even those with a structural bias towards learning nested tree (i.e., stack-RNNs). In addition, our analyses revealed primacy and recency effects in the generalization patterns of LSTM-based models, showing that these models tend to perform well on the outer- and innermost parts of a center-embedded tree structure, but poorly on its middle levels. Finally, probing the internal states of the model during the processing of sentences with nested tree structures, we found a complex encoding of grammatical agreement information (e.g. grammatical number), in which all the information for multiple words nouns was carried by a single unit. Taken together, these results indicate how neural networks may extract bounded nested tree structures, without learning a systematic recursive rule.