Pre-training Co-evolutionary Protein Representation via A Pairwise Masked Language Model
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Abstract (arXiv)
Understanding protein sequences is vital and urgent for biology, healthcare, and medicine. Labeling approaches are expensive yet time-consuming, while the amount of unlabeled data is increasing quite faster than that of the labeled data due to low-cost, high-throughput sequencing methods. In order to extract knowledge from these unlabeled data, representation learning is of significant value for protein-related tasks and has great potential for helping us learn more about protein functions and structures. The key problem in the protein sequence representation learning is to capture the co-evolutionary information reflected by the inter-residue co-variation in the sequences. Instead of leveraging multiple sequence alignment as is usually done, we propose a novel method to capture this information directly by pre-training via a dedicated language model, i.e., Pairwise Masked Language Model (PMLM). In a conventional masked language model, the masked tokens are modeled by conditioning on the unmasked tokens only, but processed independently to each other. However, our proposed PMLM takes the dependency among masked tokens into consideration, i.e., the probability of a token pair is not equal to the product of the probability of the two tokens. By applying this model, the pre-trained encoder is able to generate a better representation for protein sequences. Our result shows that the proposed method can effectively capture the inter-residue correlations and improves the performance of contact prediction by up to 9% compared to the MLM baseline under the same setting. The proposed model also significantly outperforms the MSA baseline by more than 7% on the TAPE contact prediction benchmark when pre-trained on a subset of the sequence database which the MSA is generated from, revealing the potential of the sequence pre-training method to surpass MSA based methods in general.
Key facts. Cited by 20 papers in the map (data through 7 Oct 2026). Builds on 14 of its 53 references that are in the map. First posted on arXiv 29 Oct 2021.
Builds on
14 of 53 references.
References · 14
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- 2019RoBERTa: A Robustly Optimized BERT Pretraining Approachcited by 13,266
- 2018Deep contextualized word representationscited by 2,285
- 2019Evaluating Protein Transfer Learning with TAPEcited by 229
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- 2020ProGen: Language Modeling for Protein Generationcited by 127
- 2020BERTology Meets Biology: Interpreting Attention in Protein Language Modelscited by 85
- 2019Learning protein sequence embeddings using information from structurecited by 50
- 2016Accurate De Novo Prediction of Protein Contact Map by Ultra-Deep Learning Modelcited by 31
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Citing papers per year: 2022: 6; 2023: 5; 2024: 7; 2025: 1; 2026: 1.
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- 2022Protein Representation Learning by Geometric Structure Pretrainingcited by 161
- 2024Diffusion Language Models Are Versatile Protein Learnerscited by 93
- 2024Scientific Large Language Models: A Survey on Biological & Chemical Domainscited by 80
- 2022PEER: A Comprehensive and Multi-Task Benchmark for Protein Sequence Understandingcited by 77
- 2023Structure-informed Language Models Are Protein Designerscited by 68
- 2024DPLM-2: A Multimodal Diffusion Protein Language Modelcited by 59
- 2022Protein Language Models and Structure Prediction: Connection and Progressioncited by 17
- 2024SciSafeEval: A Comprehensive Benchmark for Safety Alignment of Large Language Models in Scientific Taskscited by 17
- 2023A Systematic Study of Joint Representation Learning on Protein Sequences and Structurescited by 15
- 2023A Hierarchical Training Paradigm for Antibody Structure-sequence Co-designcited by 13
- 2023Masked Modeling for Self-supervised Representation Learning on Vision and Beyondcited by 13
- 2022A Survey on Protein Representation Learning: Retrospect and Prospectcited by 10
8 more
- 2023Retrieved Sequence Augmentation for Protein Representation Learningcited by 7
- 2025Computational Protein Science in the Era of Large Language Models (LLMs)cited by 6
- 2022Incorporating Pre-training Paradigm for Antibody Sequence-Structure Co-designcited by 3
- 2024Beyond ESM2: Graph-Enhanced Protein Sequence Modeling with Efficient Clusteringcited by 1
- 2024Long-context Protein Language Modeling Using Bidirectional Mamba with Shared Projection Layerscited by 1
- 2026CAPSUL: A Comprehensive Human Protein Benchmark for Subcellular Localizationcited by 1
- 2022Training self-supervised peptide sequence models on artificially chopped proteinscited by 0
- 2024Could Chemical LLMs benefit from Message Passingcited by 0
- 2026CAPSUL: A Comprehensive Human Protein Benchmark for Subcellular Localization19 Mar 2026
- 2025Computational Protein Science in the Era of Large Language Models (LLMs)17 Jan 2025
- 2024Long-context Protein Language Modeling Using Bidirectional Mamba with Shared Projection Layers29 Oct 2024
- 2024DPLM-2: A Multimodal Diffusion Protein Language Model17 Oct 2024
- 2024SciSafeEval: A Comprehensive Benchmark for Safety Alignment of Large Language Models in Scientific Tasks2 Oct 2024
- 2024Could Chemical LLMs benefit from Message Passing14 May 2024
- 2024Beyond ESM2: Graph-Enhanced Protein Sequence Modeling with Efficient Clustering24 Apr 2024
- 2024Diffusion Language Models Are Versatile Protein Learners28 Feb 2024
- 2024Scientific Large Language Models: A Survey on Biological & Chemical Domains26 Jan 2024
- 2023Masked Modeling for Self-supervised Representation Learning on Vision and Beyond31 Dec 2023
- 2023A Hierarchical Training Paradigm for Antibody Structure-sequence Co-design30 Oct 2023
- 2023A Systematic Study of Joint Representation Learning on Protein Sequences and Structures11 Mar 2023
8 more
- 2023Retrieved Sequence Augmentation for Protein Representation Learning24 Feb 2023
- 2023Structure-informed Language Models Are Protein Designers3 Feb 2023
- 2022A Survey on Protein Representation Learning: Retrospect and Prospect31 Dec 2022
- 2022Protein Language Models and Structure Prediction: Connection and Progression30 Nov 2022
- 2022Training self-supervised peptide sequence models on artificially chopped proteins9 Nov 2022
- 2022Incorporating Pre-training Paradigm for Antibody Sequence-Structure Co-design26 Oct 2022
- 2022PEER: A Comprehensive and Multi-Task Benchmark for Protein Sequence Understanding5 Jun 2022
- 2022Protein Representation Learning by Geometric Structure Pretraining11 Mar 2022
- 2024Diffusion Language Models Are Versatile Protein Learnerscited by 93
- 2024Diffusion Language Models Are Versatile Protein Learners28 Feb 2024
- 2022Protein Representation Learning by Geometric Structure Pretrainingcited by 161
- 2024Scientific Large Language Models: A Survey on Biological & Chemical Domainscited by 80
- 2022PEER: A Comprehensive and Multi-Task Benchmark for Protein Sequence Understandingcited by 77
- 2023Structure-informed Language Models Are Protein Designerscited by 68
- 2024DPLM-2: A Multimodal Diffusion Protein Language Modelcited by 59
- 2022Protein Language Models and Structure Prediction: Connection and Progressioncited by 17
- 2023A Systematic Study of Joint Representation Learning on Protein Sequences and Structurescited by 15
- 2023A Hierarchical Training Paradigm for Antibody Structure-sequence Co-designcited by 13
- 2023Masked Modeling for Self-supervised Representation Learning on Vision and Beyondcited by 13
- 2022A Survey on Protein Representation Learning: Retrospect and Prospectcited by 10
- 2023Retrieved Sequence Augmentation for Protein Representation Learningcited by 7
- 2025Computational Protein Science in the Era of Large Language Models (LLMs)cited by 6
12 most cited of 17 All 17 (JSON)
- 2026CAPSUL: A Comprehensive Human Protein Benchmark for Subcellular Localization19 Mar 2026
- 2025Computational Protein Science in the Era of Large Language Models (LLMs)17 Jan 2025
- 2024DPLM-2: A Multimodal Diffusion Protein Language Model17 Oct 2024
- 2024Could Chemical LLMs benefit from Message Passing14 May 2024
- 2024Beyond ESM2: Graph-Enhanced Protein Sequence Modeling with Efficient Clustering24 Apr 2024
- 2024Scientific Large Language Models: A Survey on Biological & Chemical Domains26 Jan 2024
- 2023Masked Modeling for Self-supervised Representation Learning on Vision and Beyond31 Dec 2023
- 2023A Hierarchical Training Paradigm for Antibody Structure-sequence Co-design30 Oct 2023
- 2023A Systematic Study of Joint Representation Learning on Protein Sequences and Structures11 Mar 2023
- 2023Retrieved Sequence Augmentation for Protein Representation Learning24 Feb 2023
- 2023Structure-informed Language Models Are Protein Designers3 Feb 2023
- 2022A Survey on Protein Representation Learning: Retrospect and Prospect31 Dec 2022
12 newest of 17 All 17 (JSON)