Jean-Rémi King

Juliette Millet, Charlotte Caucheteux, Pierre Orhan et al.

Several deep neural networks have recently been shown to generate activations similar to those of the brain in response to the same input. These algorithms, however, remain largely implausible: they require (1) extraordinarily large amounts of data, (2) unobtainable supervised labels, (3) textual rather than raw sensory input, and / or (4) implausibly large memory (e.g. thousands of contextual words). These elements highlight the need to identify algorithms that, under these limitations, would suffice to account for both behavioral and brain responses. Focusing on the issue of speech processing, we here hypothesize that self-supervised algorithms trained on the raw waveform constitute a promising candidate. Specifically, we compare a recent self-supervised architecture, Wav2Vec 2.0, to the brain activity of 412 English, French, and Mandarin individuals recorded with functional Magnetic Resonance Imaging (fMRI), while they listened to ~1h of audio books. Our results are four-fold. First, we show that this algorithm learns brain-like representations with as little as 600 hours of unlabelled speech -- a quantity comparable to what infants can be exposed to during language acquisition. Second, its functional hierarchy aligns with the cortical hierarchy of speech processing. Third, different training regimes reveal a functional specialization akin to the cortex: Wav2Vec 2.0 learns sound-generic, speech-specific and language-specific representations similar to those of the prefrontal and temporal cortices. Fourth, we confirm the similarity of this specialization with the behavior of 386 additional participants. These elements, resulting from the largest neuroimaging benchmark to date, show how self-supervised learning can account for a rich organization of speech processing in the brain, and thus delineate a path to identify the laws of language acquisition which shape the human brain.

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.

Laura Gwilliams, Graham Flick, Alec Marantz et al.

The "MEG-MASC" dataset provides a curated set of raw magnetoencephalography (MEG) recordings of 27 English speakers who listened to two hours of naturalistic stories. Each participant performed two identical sessions, involving listening to four fictional stories from the Manually Annotated Sub-Corpus (MASC) intermixed with random word lists and comprehension questions. We time-stamp the onset and offset of each word and phoneme in the metadata of the recording, and organize the dataset according to the 'Brain Imaging Data Structure' (BIDS). This data collection provides a suitable benchmark to large-scale encoding and decoding analyses of temporally-resolved brain responses to speech. We provide the Python code to replicate several validations analyses of the MEG evoked related fields such as the temporal decoding of phonetic features and word frequency. All code and MEG, audio and text data are publicly available to keep with best practices in transparent and reproducible research.

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.

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, 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.

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.

Bruno Aristimunha, Dung Truong, Pierre Guetschel et al.

Current electroencephalogram (EEG) decoding models are typically trained on small numbers of subjects performing a single task. Here, we introduce a large-scale, code-submission-based competition comprising two challenges. First, the Transfer Challenge asks participants to build and test a model that can zero-shot decode new tasks and new subjects from their EEG data. Second, the Psychopathology factor prediction Challenge asks participants to infer subject measures of mental health from EEG data. For this, we use an unprecedented, multi-terabyte dataset of high-density EEG signals (128 channels) recorded from over 3,000 child to young adult subjects engaged in multiple active and passive tasks. We provide several tunable neural network baselines for each of these two challenges, including a simple network and demographic-based regression models. Developing models that generalise across tasks and individuals will pave the way for ML network architectures capable of adapting to EEG data collected from diverse tasks and individuals. Similarly, predicting mental health-relevant personality trait values from EEG might identify objective biomarkers useful for clinical diagnosis and design of personalised treatment for psychological conditions. Ultimately, the advances spurred by this challenge could contribute to the development of computational psychiatry and useful neurotechnology, and contribute to breakthroughs in both fundamental neuroscience and applied clinical research.

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-Remi King

Deep learning has recently made remarkable progress in natural language processing. Yet, the resulting algorithms remain far from competing with the language abilities of the human brain. Predictive coding theory offers a potential explanation to this discrepancy: while deep language algorithms are optimized to predict adjacent words, the human brain would be tuned to make long-range and hierarchical predictions. To test this hypothesis, we analyze the fMRI brain signals of 304 subjects each listening to 70min of short stories. After confirming that the activations of deep language algorithms linearly map onto those of the brain, we show that enhancing these models with long-range forecast representations improves their brain-mapping. The results further reveal a hierarchy of predictions in the brain, whereby the fronto-parietal cortices forecast more abstract and more distant representations than the temporal cortices. Overall, this study strengthens predictive coding theory and suggests a critical role of long-range and hierarchical predictions in natural language 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.

Omar Chehab, Alexandre Defossez, Jean-Christophe Loiseau et al.

Understanding how the brain responds to sensory inputs is challenging: brain recordings are partial, noisy, and high dimensional; they vary across sessions and subjects and they capture highly nonlinear dynamics. These challenges have led the community to develop a variety of preprocessing and analytical (almost exclusively linear) methods, each designed to tackle one of these issues. Instead, we propose to address these challenges through a specific end-to-end deep learning architecture, trained to predict the brain responses of multiple subjects at once. We successfully test this approach on a large cohort of magnetoencephalography (MEG) recordings acquired during a one-hour reading task. Our Deep Recurrent Encoding (DRE) architecture reliably predicts MEG responses to words with a three-fold improvement over classic linear methods. To overcome the notorious issue of interpretability of deep learning, we describe a simple variable importance analysis. When applied to DRE, this method recovers the expected evoked responses to word length and word frequency. The quantitative improvement of the present deep learning approach paves the way to better understand the nonlinear dynamics of brain activity from large datasets.

Charlotte Caucheteux, Alexandre Gramfort, Jean-Remi King

The activations of language transformers like GPT-2 have been shown to linearly map onto brain activity during speech comprehension. However, the nature of these activations remains largely unknown and presumably conflate distinct linguistic classes. Here, we propose a taxonomy to factorize the high-dimensional activations of language models into four combinatorial classes: lexical, compositional, syntactic, and semantic representations. We then introduce a statistical method to decompose, through the lens of GPT-2's activations, the brain activity of 345 subjects recorded with functional magnetic resonance imaging (fMRI) during the listening of ~4.6 hours of narrated text. The results highlight two findings. First, compositional representations recruit a more widespread cortical network than lexical ones, and encompass the bilateral temporal, parietal and prefrontal cortices. Second, contrary to previous claims, syntax and semantics are not associated with separated modules, but, instead, appear to share a common and distributed neural substrate. Overall, this study introduces a versatile framework to isolate, in the brain activity, the distributed representations of linguistic constructs.

Juliette Millet, Jean-Remi King

Our ability to comprehend speech remains, to date, unrivaled by deep learning models. This feat could result from the brain's ability to fine-tune generic sound representations for speech-specific processes. To test this hypothesis, we compare i) five types of deep neural networks to ii) human brain responses elicited by spoken sentences and recorded in 102 Dutch subjects using functional Magnetic Resonance Imaging (fMRI). Each network was either trained on an acoustics scene classification, a speech-to-text task (based on Bengali, English, or Dutch), or not trained. The similarity between each model and the brain is assessed by correlating their respective activations after an optimal linear projection. The differences in brain-similarity across networks revealed three main results. First, speech representations in the brain can be accounted for by random deep networks. Second, learning to classify acoustic scenes leads deep nets to increase their brain similarity. Third, learning to process phonetically-related speech inputs (i.e., Dutch vs English) leads deep nets to reach higher levels of brain-similarity than learning to process phonetically-distant speech inputs (i.e. Dutch vs Bengali). Together, these results suggest that the human brain fine-tunes its heavily-trained auditory hierarchy to learn to process speech.

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.