Disentangling Syntax and Semantics in the Brain with Deep Networks
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Abstract (arXiv)
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.
Key facts. Cited by 39 papers in the map (data through 7 Oct 2026). Builds on 17 of its 67 references that are in the map. First posted on arXiv 2 Mar 2021.
Builds on
17 of 67 references.
References · 17
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- 2020Language Models are Few-Shot Learnerscited by 32,199
- 2019RoBERTa: A Robustly Optimized BERT Pretraining Approachcited by 13,266
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- 2019Cross-lingual Language Model Pretrainingcited by 1,088
- 2019Compositionality decomposed: how do neural networks generalise?cited by 264
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- 2019Interpreting and improving natural-language processing (in machines) with natural language-processing (in the brain)cited by 128
- 2020Syntactic Structure from Deep Learningcited by 99
- 2019The emergence of number and syntax units in LSTM language modelscited by 92
5 more
- 2019Linguistic generalization and compositionality in modern artificial neural networkscited by 58
- 2020Efficient Second-Order TreeCRF for Neural Dependency Parsingcited by 48
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Citing papers per year: 2021: 1; 2022: 5; 2023: 8; 2024: 11; 2025: 7; 2026: 7.
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- 2022Toward a realistic model of speech processing in the brain with self-supervised learningcited by 63
- 2023Explaining black box text modules in natural language with language modelscited by 61
- 2023Emergent Modularity in Pre-trained Transformerscited by 37
- 2024Contextual Feature Extraction Hierarchies Converge in Large Language Models and the Braincited by 32
- 2022Neural Language Taskonomy: Which NLP Tasks are the most Predictive of fMRI Brain Activity?cited by 27
- 2022Neural Language Models are not Born Equal to Fit Brain Data, but Training Helpscited by 26
- 2024Decoding Probing: Revealing Internal Linguistic Structures in Neural Language Models using Minimal Pairscited by 24
- 2024A polar coordinate system represents syntax in large language modelscited by 24
- 2023Deep Neural Networks and Brain Alignment: Brain Encoding and Decoding (Survey)cited by 20
- 2023Information-Restricted Neural Language Models Reveal Different Brain Regions' Sensitivity to Semantics, Syntax and Contextcited by 13
- 2024Crafting Interpretable Embeddings by Asking LLMs Questionscited by 11
- 2021Long-range and hierarchical language predictions in brains and algorithmscited by 10
27 more
- 2024What Are Large Language Models Mapping to in the Brain? A Case Against Over-Reliance on Brain Scorescited by 10
- 2023Coupling Artificial Neurons in BERT and Biological Neurons in the Human Braincited by 9
- 2023Mapping Brains with Language Models: A Surveycited by 8
- 2024fMRI predictors based on language models of increasing complexity recover brain left lateralizationcited by 8
- 2022Connecting Neural Response measurements & Computational Models of language: a non-comprehensive guidecited by 6
- 2025Decoupling Content and Expression: Two-Dimensional Detection of AI-Generated Textcited by 6
- 2026Differential syntactic and semantic encoding in LLMscited by 6
- 2024Multilevel Interpretability Of Artificial Neural Networks: Leveraging Framework And Methods From Neurosciencecited by 5
- 2024Computational Models to Study Language Processing in the Human Brain: A Surveycited by 4
- 2025Explanations of Large Language Models Explain Language Representations in the Braincited by 4
- 2025Probing Syntax in Large Language Models: Successes and Remaining Challengescited by 4
- 2023Deep Learning Models to Study Sentence Comprehension in the Human Braincited by 3
- 2022MEG-MASC: a high-quality magneto-encephalography dataset for evaluating natural speech processingcited by 2
- 2024Interpretable Next-token Prediction via the Generalized Induction Headcited by 2
- 2025Hierarchical Frequency Tagging Probe (HFTP): A Unified Approach to Investigate Syntactic Structure Representations in Large Language Models and the Human Braincited by 2
- 2025Emergence and Localisation of Semantic Role Circuits in LLMscited by 2
- 2026Sparse Autoencoders Map Brain-LLM Alignment onto Cortical Semantic Topographycited by 2
- 2023Syntactic Structure Processing in the Brain while Listeningcited by 1
- 2024Decoding Linguistic Representations of Human Braincited by 1
- 2025Improve Language Model and Brain Alignment via Associative Memorycited by 1
- 2024Investigating the Timescales of Language Processing with EEG and Language Modelscited by 0
- 2025Testing Transformer Learnability on the Arithmetic Sequence of Rooted Treescited by 0
- 2026Left-right asymmetry in predicting brain activity from LLMs' representations emerges with their formal linguistic competencecited by 0
- 2026Computational Lesions in Multilingual Language Models Separate Shared and Language-specific Brain Alignmentcited by 0
- 2026Position: AI for Science Should Treat Measurement-to-Dataset Pipelines as Inference Componentscited by 0
- 2026Triospect: A Three-Dimensional Framework for Robust Statistical AI-Generated Text Detection Against Diverse Attackscited by 0
- 2026Syntax vs. Semantics: How Transformers Learn Deep Dependenciescited by 0
- 2026Triospect: A Three-Dimensional Framework for Robust Statistical AI-Generated Text Detection Against Diverse Attacks30 Jun 2026
- 2026Syntax vs. Semantics: How Transformers Learn Deep Dependencies26 Jun 2026
- 2026Position: AI for Science Should Treat Measurement-to-Dataset Pipelines as Inference Components23 May 2026
- 2026Sparse Autoencoders Map Brain-LLM Alignment onto Cortical Semantic Topography21 May 2026
- 2026Computational Lesions in Multilingual Language Models Separate Shared and Language-specific Brain Alignment12 Apr 2026
- 2026Left-right asymmetry in predicting brain activity from LLMs' representations emerges with their formal linguistic competence13 Feb 2026
- 2026Differential syntactic and semantic encoding in LLMs8 Jan 2026
- 2025Testing Transformer Learnability on the Arithmetic Sequence of Rooted Trees1 Dec 2025
- 2025Emergence and Localisation of Semantic Role Circuits in LLMs25 Nov 2025
- 2025Hierarchical Frequency Tagging Probe (HFTP): A Unified Approach to Investigate Syntactic Structure Representations in Large Language Models and the Human Brain15 Oct 2025
- 2025Probing Syntax in Large Language Models: Successes and Remaining Challenges5 Aug 2025
- 2025Improve Language Model and Brain Alignment via Associative Memory20 May 2025
27 more
- 2025Decoupling Content and Expression: Two-Dimensional Detection of AI-Generated Text1 Mar 2025
- 2025Explanations of Large Language Models Explain Language Representations in the Brain20 Feb 2025
- 2024A polar coordinate system represents syntax in large language models7 Dec 2024
- 2024Interpretable Next-token Prediction via the Generalized Induction Head31 Oct 2024
- 2024Multilevel Interpretability Of Artificial Neural Networks: Leveraging Framework And Methods From Neuroscience22 Aug 2024
- 2024Decoding Linguistic Representations of Human Brain30 Jul 2024
- 2024Investigating the Timescales of Language Processing with EEG and Language Models28 Jun 2024
- 2024What Are Large Language Models Mapping to in the Brain? A Case Against Over-Reliance on Brain Scores3 Jun 2024
- 2024fMRI predictors based on language models of increasing complexity recover brain left lateralization28 May 2024
- 2024Crafting Interpretable Embeddings by Asking LLMs Questions26 May 2024
- 2024Decoding Probing: Revealing Internal Linguistic Structures in Neural Language Models using Minimal Pairs26 Mar 2024
- 2024Computational Models to Study Language Processing in the Human Brain: A Survey20 Mar 2024
- 2024Contextual Feature Extraction Hierarchies Converge in Large Language Models and the Brain31 Jan 2024
- 2023Deep Neural Networks and Brain Alignment: Brain Encoding and Decoding (Survey)17 Jul 2023
- 2023Mapping Brains with Language Models: A Survey8 Jun 2023
- 2023Emergent Modularity in Pre-trained Transformers28 May 2023
- 2023Explaining black box text modules in natural language with language models17 May 2023
- 2023Coupling Artificial Neurons in BERT and Biological Neurons in the Human Brain27 Mar 2023
- 2023Information-Restricted Neural Language Models Reveal Different Brain Regions' Sensitivity to Semantics, Syntax and Context28 Feb 2023
- 2023Syntactic Structure Processing in the Brain while Listening16 Feb 2023
- 2023Deep Learning Models to Study Sentence Comprehension in the Human Brain16 Jan 2023
- 2022MEG-MASC: a high-quality magneto-encephalography dataset for evaluating natural speech processing26 Jul 2022
- 2022Neural Language Models are not Born Equal to Fit Brain Data, but Training Helps7 Jul 2022
- 2022Toward a realistic model of speech processing in the brain with self-supervised learning3 Jun 2022
- 2022Neural Language Taskonomy: Which NLP Tasks are the most Predictive of fMRI Brain Activity?3 May 2022
- 2022Connecting Neural Response measurements & Computational Models of language: a non-comprehensive guide10 Mar 2022
- 2021Long-range and hierarchical language predictions in brains and algorithms28 Nov 2021
- 2022Toward a realistic model of speech processing in the brain with self-supervised learningcited by 63
- 2023Explaining black box text modules in natural language with language modelscited by 61
- 2023Emergent Modularity in Pre-trained Transformerscited by 37
- 2024Contextual Feature Extraction Hierarchies Converge in Large Language Models and the Braincited by 32
- 2022Neural Language Taskonomy: Which NLP Tasks are the most Predictive of fMRI Brain Activity?cited by 27
- 2022Neural Language Models are not Born Equal to Fit Brain Data, but Training Helpscited by 26
- 2024Decoding Probing: Revealing Internal Linguistic Structures in Neural Language Models using Minimal Pairscited by 24
- 2024A polar coordinate system represents syntax in large language modelscited by 24
- 2023Deep Neural Networks and Brain Alignment: Brain Encoding and Decoding (Survey)cited by 20
- 2023Information-Restricted Neural Language Models Reveal Different Brain Regions' Sensitivity to Semantics, Syntax and Contextcited by 13
- 2024Crafting Interpretable Embeddings by Asking LLMs Questionscited by 11
- 2021Long-range and hierarchical language predictions in brains and algorithmscited by 10
12 most cited of 38 All 38 (JSON)
- 2026Triospect: A Three-Dimensional Framework for Robust Statistical AI-Generated Text Detection Against Diverse Attacks30 Jun 2026
- 2026Syntax vs. Semantics: How Transformers Learn Deep Dependencies26 Jun 2026
- 2026Position: AI for Science Should Treat Measurement-to-Dataset Pipelines as Inference Components23 May 2026
- 2026Computational Lesions in Multilingual Language Models Separate Shared and Language-specific Brain Alignment12 Apr 2026
- 2026Left-right asymmetry in predicting brain activity from LLMs' representations emerges with their formal linguistic competence13 Feb 2026
- 2026Differential syntactic and semantic encoding in LLMs8 Jan 2026
- 2025Testing Transformer Learnability on the Arithmetic Sequence of Rooted Trees1 Dec 2025
- 2025Emergence and Localisation of Semantic Role Circuits in LLMs25 Nov 2025
- 2025Hierarchical Frequency Tagging Probe (HFTP): A Unified Approach to Investigate Syntactic Structure Representations in Large Language Models and the Human Brain15 Oct 2025
- 2025Probing Syntax in Large Language Models: Successes and Remaining Challenges5 Aug 2025
- 2025Improve Language Model and Brain Alignment via Associative Memory20 May 2025
- 2025Decoupling Content and Expression: Two-Dimensional Detection of AI-Generated Text1 Mar 2025
12 newest of 38 All 38 (JSON)