Greta Tuckute

Jian Li, Greta Tuckute, Evelina Fedorenko et al.

Surface-based cortical registration is an important topic in medical image analysis and facilitates many downstream applications. Current approaches for cortical registration are mainly driven by geometric features, such as sulcal depth and curvature, and often assume that registration of folding patterns leads to alignment of brain function. However, functional variability of anatomically corresponding areas across subjects has been widely reported, particularly in higher-order cognitive areas. In this work, we present JOSA, a novel cortical registration framework that jointly models the mismatch between geometry and function while simultaneously learning an unbiased population-specific atlas. Using a semi-supervised training strategy, JOSA achieves superior registration performance in both geometry and function to the state-of-the-art methods but without requiring functional data at inference. This learning framework can be extended to any auxiliary data to guide spherical registration that is available during training but is difficult or impossible to obtain during inference, such as parcellations, architectonic identity, transcriptomic information, and molecular profiles. By recognizing the mismatch between geometry and function, JOSA provides new insights into the future development of registration methods using joint analysis of the brain structure and function.

Jian Li, Greta Tuckute, Evelina Fedorenko et al.

Brain surface-based image registration, an important component of brain image analysis, establishes spatial correspondence between cortical surfaces. Existing iterative and learning-based approaches focus on accurate registration of folding patterns of the cerebral cortex, and assume that geometry predicts function and thus functional areas will also be well aligned. However, structure/functional variability of anatomically corresponding areas across subjects has been widely reported. In this work, we introduce a learning-based cortical registration framework, JOSA, which jointly aligns folding patterns and functional maps while simultaneously learning an optimal atlas. We demonstrate that JOSA can substantially improve registration performance in both anatomical and functional domains over existing methods. By employing a semi-supervised training strategy, the proposed framework obviates the need for functional data during inference, enabling its use in broad neuroscientific domains where functional data may not be observed. The source code of JOSA will be released to the public at https://voxelmorph.net.

Daria Kryvosheieva, Andrea de Varda, Evelina Fedorenko et al.

Large language models (LLMs) can reliably distinguish grammatical from ungrammatical sentences, but how grammatical knowledge is represented within the models remains an open question. We investigate whether different syntactic phenomena recruit shared or distinct components in LLMs. Using a functional localization approach inspired by cognitive neuroscience, we identify the LLM units most responsive to 67 English syntactic phenomena in seven open-weight models. These units are consistently recruited across sentences containing the phenomena and causally support the models' syntactic performance. Critically, different types of syntactic agreement (e.g., subject-verb, anaphor, determiner-noun) recruit overlapping sets of units, suggesting that agreement constitutes a meaningful functional category for LLMs. This pattern holds in English, Russian, and Chinese; and further, in a cross-lingual analysis of 57 diverse languages, structurally more similar languages share more units for subject-verb agreement. Taken together, these findings reveal that syntactic agreement-a critical marker of syntactic dependencies-constitutes a meaningful category within LLMs' representational spaces.

Ekdeep Singh Lubana, Can Rager, Sai Sumedh R. Hindupur et al.

Recovering meaningful concepts from language model activations is a central aim of interpretability. While existing feature extraction methods aim to identify concepts that are independent directions, it is unclear if this assumption can capture the rich temporal structure of language. Specifically, via a Bayesian lens, we demonstrate that Sparse Autoencoders (SAEs) impose priors that assume independence of concepts across time, implying stationarity. Meanwhile, language model representations exhibit rich temporal dynamics, including systematic growth in conceptual dimensionality, context-dependent correlations, and pronounced non-stationarity, in direct conflict with the priors of SAEs. Taking inspiration from computational neuroscience, we introduce a new interpretability objective -- Temporal Feature Analysis -- which possesses a temporal inductive bias to decompose representations at a given time into two parts: a predictable component, which can be inferred from the context, and a residual component, which captures novel information unexplained by the context. Temporal Feature Analyzers correctly parse garden path sentences, identify event boundaries, and more broadly delineate abstract, slow-moving information from novel, fast-moving information, while existing SAEs show significant pitfalls in all the above tasks. Overall, our results underscore the need for inductive biases that match the data in designing robust interpretability tools.

Badr AlKhamissi, Greta Tuckute, Antoine Bosselut et al.

Large language models (LLMs) exhibit remarkable capabilities on not just language tasks, but also various tasks that are not linguistic in nature, such as logical reasoning and social inference. In the human brain, neuroscience has identified a core language system that selectively and causally supports language processing. We here ask whether similar specialization for language emerges in LLMs. We identify language-selective units within 18 popular LLMs, using the same localization approach that is used in neuroscience. We then establish the causal role of these units by demonstrating that ablating LLM language-selective units -- but not random units -- leads to drastic deficits in language tasks. Correspondingly, language-selective LLM units are more aligned to brain recordings from the human language system than random units. Finally, we investigate whether our localization method extends to other cognitive domains: while we find specialized networks in some LLMs for reasoning and social capabilities, there are substantial differences among models. These findings provide functional and causal evidence for specialization in large language models, and highlight parallels with the functional organization in the brain.

Badr AlKhamissi, Greta Tuckute, Yingtian Tang et al.

Large language models (LLMs) exhibit remarkable similarity to neural activity in the human language network. However, the key properties of language shaping brain-like representations, and their evolution during training as a function of different tasks remain unclear. We here benchmark 34 training checkpoints spanning 300B tokens across 8 different model sizes to analyze how brain alignment relates to linguistic competence. Specifically, we find that brain alignment tracks the development of formal linguistic competence -- i.e., knowledge of linguistic rules -- more closely than functional linguistic competence. While functional competence, which involves world knowledge and reasoning, continues to develop throughout training, its relationship with brain alignment is weaker, suggesting that the human language network primarily encodes formal linguistic structure rather than broader cognitive functions. We further show that model size is not a reliable predictor of brain alignment when controlling for feature size and find that the correlation between next-word prediction, behavioral alignment and brain alignment fades once models surpass human language proficiency. Finally, using the largest set of rigorous neural language benchmarks to date, we show that language brain alignment benchmarks remain unsaturated, highlighting opportunities for improving future models. Taken together, our findings suggest that the human language network is best modeled by formal, rather than functional, aspects of language.

Taha Binhuraib, Greta Tuckute, Nicholas Blauch

Spatial functional organization is a hallmark of biological brains: neurons are arranged topographically according to their response properties, at multiple scales. In contrast, representations within most machine learning models lack spatial biases, instead manifesting as disorganized vector spaces that are difficult to visualize and interpret. Here, we propose a novel form of self-attention that turns Transformers into "Topoformers" with topographic organization. We introduce spatial querying - where keys and queries are arranged on 2D grids, and local pools of queries are associated with a given key - and spatial reweighting, where we convert the standard fully connected layer of self-attention into a locally connected layer. We first demonstrate the feasibility of our approach by training a 1-layer Topoformer on a sentiment classification task. Training with spatial querying encourages topographic organization in the queries and keys, and spatial reweighting separately encourages topographic organization in the values and self-attention outputs. We then apply the Topoformer motifs at scale, training a BERT architecture with a masked language modeling objective. We find that the topographic variant performs on par with a non-topographic control model on NLP benchmarks, yet produces interpretable topographic organization as evaluated via eight linguistic test suites. Finally, analyzing an fMRI dataset of human brain responses to a large set of naturalistic sentences, we demonstrate alignment between low-dimensional topographic variability in the Topoformer model and human brain language network. Scaling up Topoformers further holds promise for greater interpretability in NLP research, and for more accurate models of the organization of linguistic information in the human brain.

Lukas Wolf, Greta Tuckute, Klemen Kotar et al.

Training on multiple modalities of input can augment the capabilities of a language model. Here, we ask whether such a training regime can improve the quality and efficiency of these systems as well. We focus on text--audio and introduce Whisbert, which is inspired by the text--image approach of FLAVA (Singh et al., 2022). In accordance with Babylm guidelines (Warstadt et al., 2023), we pretrain Whisbert on a dataset comprising only 100 million words plus their corresponding speech from the word-aligned version of the People's Speech dataset (Galvez et al., 2021). To assess the impact of multimodality, we compare versions of the model that are trained on text only and on both audio and text simultaneously. We find that while Whisbert is able to perform well on multimodal masked modeling and surpasses the Babylm baselines in most benchmark tasks, it struggles to optimize its complex objective and outperform its text-only Whisbert baseline.

Badr AlKhamissi, C. Nicolò De Sabbata, Greta Tuckute et al.

Human cognitive behavior arises from the interaction of specialized brain networks dedicated to distinct functions, such as language, logic, and social reasoning. Inspired by this organization, we propose Mixture of Cognitive Reasoners (MiCRo): a modular, transformer-based architecture post-trained with a curriculum that induces functional specialization across experts. Concretely, we partition the layers of a pretrained language model into four expert modules aligned with well-studied cognitive networks in the human brain. MiCRo offers three key advantages over standard language models. (1) The specialized experts are interpretable and causally meaningful -- ablating a module causes substantial drops on benchmarks requiring its specialized domain. (2) MiCRo's behavior can be dynamically steered at inference time by routing tokens to particular experts (e.g., favoring social over logical reasoning), enabling fine-grained control over outputs. (3) MiCRo outperforms or matches comparable baselines on both machine-learning reasoning benchmarks (e.g., GSM8K, BBH) and alignment to human behavior (CogBench), while maintaining interpretability. Taken together, cognitively grounded functional specialization yields models that are both more human-like and more human-interpretable.

Maria Ryskina, Greta Tuckute, Alexander Fung et al. · mit

Cognitive science and neuroscience have long faced the challenge of disentangling representations of language from representations of conceptual meaning. As the same problem arises in today's language models (LMs), we investigate the relationship between LM--brain alignment and two neural metrics: (1) the level of brain activation during processing of sentences, targeting linguistic processing, and (2) a novel measure of meaning consistency across input modalities, which quantifies how consistently a brain region responds to the same concept across paradigms (sentence, word cloud, image) using an fMRI dataset (Pereira et al., 2018). Our experiments show that both language-only and language-vision models predict the signal better in more meaning-consistent areas of the brain, even when these areas are not strongly sensitive to language processing, suggesting that LMs might internally represent cross-modal conceptual meaning.

Shreya Saha, Shurui Li, Greta Tuckute et al.

The human language system represents both linguistic forms and meanings, but the abstractness of the meaning representations remains debated. Here, we searched for abstract representations of meaning in the language cortex by modeling neural responses to sentences using representations from vision and language models. When we generate images corresponding to sentences and extract vision model embeddings, we find that aggregating across multiple generated images yields increasingly accurate predictions of language cortex responses, sometimes rivaling large language models. Similarly, averaging embeddings across multiple paraphrases of a sentence improves prediction accuracy compared to any single paraphrase. Enriching paraphrases with contextual details that may be implicit (e.g., augmenting "I had a pancake" to include details like "maple syrup") further increases prediction accuracy, even surpassing predictions based on the embedding of the original sentence, suggesting that the language system maintains richer and broader semantic representations than language models. Together, these results demonstrate the existence of highly abstract, form-independent meaning representations within the language cortex.

Greta Tuckute, Klemen Kotar, Evelina Fedorenko et al.

We introduce AuriStream, a biologically inspired model for encoding speech via a two-stage framework inspired by the human auditory processing hierarchy. The first stage transforms raw audio into a time-frequency representation based on the human cochlea, from which we extract discrete \textbf{cochlear tokens}. The second stage applies an autoregressive sequence model over the cochlear tokens. AuriStream learns meaningful phoneme and word representations, and state-of-the-art lexical semantics. AuriStream shows competitive performance on diverse downstream SUPERB speech tasks. Complementing AuriStream's strong representational capabilities, it generates continuations of audio which can be visualized in a spectrogram space and decoded back into audio, providing insights into the model's predictions. In summary, we present a two-stage framework for speech representation learning to advance the development of more human-like models that efficiently handle a range of speech-based tasks.

Klemen Kotar, Greta Tuckute

Biological neural networks are shaped both by evolution across generations and by individual learning within an organism's lifetime, whereas standard artificial neural networks undergo a single, large training procedure without inherited constraints. In this preliminary work, we propose a framework that incorporates this crucial generational dimension - an "outer loop" of evolution that shapes the "inner loop" of learning - so that artificial networks better mirror the effects of evolution and individual learning in biological organisms. Focusing on language, we train a model that inherits a "model connectome" from the outer evolution loop before exposing it to a developmental-scale corpus of 100M tokens. Compared with two closely matched control models, we show that the connectome model performs better or on par on natural language processing tasks as well as alignment to human behavior and brain data. These findings suggest that a model connectome serves as an efficient prior for learning in low-data regimes - narrowing the gap between single-generation artificial models and biologically evolved neural networks.

Badr AlKhamissi, Greta Tuckute, Antoine Bosselut et al.

Large Language Models (LLMs) have been shown to be effective models of the human language system, with some models predicting most explainable variance of brain activity in current datasets. Even in untrained models, the representations induced by architectural priors can exhibit reasonable alignment to brain data. In this work, we investigate the key architectural components driving the surprising alignment of untrained models. To estimate LLM-to-brain similarity, we first select language-selective units within an LLM, similar to how neuroscientists identify the language network in the human brain. We then benchmark the brain alignment of these LLM units across five different brain recording datasets. By isolating critical components of the Transformer architecture, we identify tokenization strategy and multihead attention as the two major components driving brain alignment. A simple form of recurrence further improves alignment. We further demonstrate this quantitative brain alignment of our model by reproducing landmark studies in the language neuroscience field, showing that localized model units -- just like language voxels measured empirically in the human brain -- discriminate more reliably between lexical than syntactic differences, and exhibit similar response profiles under the same experimental conditions. Finally, we demonstrate the utility of our model's representations for language modeling, achieving improved sample and parameter efficiency over comparable architectures. Our model's estimates of surprisal sets a new state-of-the-art in the behavioral alignment to human reading times. Taken together, we propose a highly brain- and behaviorally-aligned model that conceptualizes the human language system as an untrained shallow feature encoder, with structural priors, combined with a trained decoder to achieve efficient and performant language processing.