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ProGen2: Exploring the boundaries of protein language models
Open accessAbstract
Attention-based models trained on protein sequences have demonstrated incredible success at classification and generation tasks relevant for artificial-intelligence-driven protein design. However, we lack a sufficient understanding of how very large-scale models and data play a role in effective protein model development. We introduce a suite of protein language models, named ProGen2, that are scaled up to 6.4B parameters and trained on different sequence datasets drawn from over a billion proteins from genomic, metagenomic, and immune repertoire databases. ProGen2 models show state-of-the-art performance in capturing the distribution of observed evolutionary sequences, generating novel viable sequences, and predicting protein fitness without additional fine-tuning. As large model sizes and raw numbers of protein sequences continue to become more widely accessible, our results suggest that a growing emphasis needs to be placed on the data distribution provided to a protein sequence model. Our models and code are open sourced for widespread adoption in protein engineering. A record of this paper's Transparent Peer Review process is included in the supplemental information.
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“Evo also reaches competitive performance with leading protein-specific language models (41, 46–48) (Fig. 2B).”
From Sequence modeling and design from molecular to genome scale with Evo (2024) · cites this paper as [48]
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Vaswani, A., Shazeer, N., Parmar, N., Uszkoreit, J., Jones, L., Gomez, A. N., Kaiser, L., & Polosukhin, I. (2017) · arXiv · doi:10.48550/arxiv.1706.03762 · Cited by 120,404
Adam: A Method for Stochastic Optimization
Kingma, D. P., & Ba, J. (2014) · arXiv · doi:10.48550/arxiv.1412.6980 · Cited by 98,946
Jakubův, J., Chvalovský, K., Goertzel, Z., et al. (2023) · DROPS (Schloss Dagstuhl – Leibniz Center for Informatics) · doi:10.4230/lipics.itp.2023.19 · Cited by 76,648
Berman, H. M. (2000) · Nucleic Acids Research · doi:10.1093/nar/28.1.235 · Cited by 40,196
Language Models are Few-Shot Learners
Brown, T. B., Mann, B., Ryder, N., Subbiah, M., Kaplan, J., Dhariwal, P., Neelakantan, A., Shyam, P., Sastry, G., Askell, A., et al. (2020) · arXiv · doi:10.48550/arxiv.2005.14165 · Cited by 23,372
Neural Machine Translation by Jointly Learning to Align and Translate
Bahdanau, D., Cho, K., & Bengio, Y. (2014) · arXiv · doi:10.48550/arxiv.1409.0473 · Cited by 16,213
Scaling Laws for Neural Language Models
Kaplan, J., McCandlish, S., Henighan, T., Brown, T. B., Chess, B., Child, R., Gray, S., Radford, A., Wu, J., & Amodei, D. (2020) · arXiv · doi:10.48550/arxiv.2001.08361 · Cited by 9,550
UniProt: a worldwide hub of protein knowledge
The UniProt Consortium (2019) · Nucleic Acids Research · doi:10.1093/nar/gky1049 · Cited by 7,698
Evolutionary-scale prediction of atomic level protein structure with a language model
Lin, Z., Akin, H., Rao, R., Hie, B., Zhu, Z., Lu, W., Smetanin, N., Verkuil, R., Kabeli, O., Shmueli, Y., et al. (2022) · doi:10.1101/2022.07.20.500902 · Cited by 5,990
MMseqs2 enables sensitive protein sequence searching for the analysis of massive data sets
Steinegger, M., & Söding, J. (2017) · Nature Biotechnology · doi:10.1038/nbt.3988 · Cited by 4,840
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Items 301–400 of 496
Protein Language Models Expose Viral Immune Mimicry
Ofer, D., & Linial, M. (2025) · Viruses · doi:10.3390/v17091199 · Cited by 5
Zhao, L., Wang, Y., Jiang, J. H., Pan, H., Wang, L., Ma, S., & Zhang, L. (2025) · Microbiology Spectrum · doi:10.1128/spectrum.01504-25 · Cited by 3
Fan, S., Chen, H., Yue, Y., Yan, K., Zou, S., Jin, Y., & Yang, Z. (2025) · Molecular Catalysis · doi:10.1016/j.mcat.2025.115515 · Cited by 2
AbTune: layer-wise selective fine-tuning of protein language models for antibodies
Xu, X., & Bonvin, A. M. J. J. (2026) · Briefings in Bioinformatics · doi:10.1093/bib/bbag374 · Cited by 2
Yang, F., Xu, R., Huang, H., Lu, J., Feng, J., Sun, X., Li, J., Du, G., & Kang, Z. (2026) · Carbohydrate Polymer Technologies and Applications · doi:10.1016/j.carpta.2026.101087 · Cited by 2
Learnable protein representations in computational biology for predicting drug-target affinity
Kumar, R., Romano, J. D., & Ritchie, M. D. (2026) · Journal of Cheminformatics · doi:10.1186/s13321-025-01145-7 · Cited by 2
Deep Learning Meets Synthetic Biology: Applications Across Molecular, Cellular, and Metabolic Scales
Zhao, Y., Yang, C., Xue, B., Liu, H., Wang, S., & Su, H. (2026) · Green Carbon · doi:10.1016/j.greenca.2025.11.003 · Cited by 2
Decoding viral evolution through integrative bioinformatics: From genomes to global health
木村, 龍., Hayashi, Y., Fujimoto-Sato, Y., Ishii, H., Suzuki, Y., Katayama, K., Mizukoshi, F., Ryo, A., & Kimura, H. (2026) · Virology · doi:10.1016/j.virol.2026.110920 · Cited by 2
Rashid, Z., Ahmed, H., Saleem, K., & Siddiqui, M. R. U. (2026) · Probiotics and Antimicrobial Proteins · doi:10.1007/s12602-026-11022-w · Cited by 2
Xiao, L., Li, J., Chen, Y., Qiu, C., Wang, Q., Liao, X., & Chen, J. (2026) · Artificial Intelligence Chemistry · doi:10.1016/j.aichem.2026.100119 · Cited by 2
Durumeric, A. E. P., McCarty, S., Smith, J., Köhler, J., Elez, K., Raich, L., Suriana, P., & Sztain, T. (2026) · Journal of Chemical Information and Modeling · doi:10.1021/acs.jcim.6c00332 · Cited by 2
Computational identification of antigen-specific T cell groups through generative epitope modeling
Ma, M., Tu, W., Vasquez-Rios, C., & Ding, J. (2026) · iScience · doi:10.1016/j.isci.2026.116505 · Cited by 2
Generative artificial intelligence for enzyme design and biocatalysis
Middendorf, L., & Ferruz, N. (2026) · Current Opinion in Chemical Biology · doi:10.1016/j.cbpa.2026.102728 · Cited by 3
A world model of the virtual cell
Xing, E. P., & Song, L. (2026) · Cell · doi:10.1016/j.cell.2026.08.042 · Cited by 2
Range-limited Heaps' law for functional DNA words in the human genome.
Li, W.-T., Almirantis, Y., & Provata, A. (2024) · Journal of Theoretical Biology · doi:10.1016/j.jtbi.2024.111878 · Cited by 4
An expandable synthetic library of human paired antibody sequences
Marinov, T. M., Wasdin, P. T., Jordaan, G., Janke, A. K., Abu-Shmais, A. A., & Georgiev, I. S. (2025) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1012932 · Cited by 2
AI expands the repertoire of CRISPR-associated proteins for genome editing
Notin, P. (2025) · Nature · doi:10.1038/d41586-025-02135-3 · Cited by 2
Using protein blocks to build custom fragment libraries from protein structures
Dhingra, S., Télétchéa, S., Sowdhamini, R., Sanejouand, Y., de Brevern, A. G., Cadet, F., & Offmann, B. (2025) · Biochimie · doi:10.1016/j.biochi.2025.08.011 · Cited by 2
Engel, R., & Park, G. (2024) · Workshop on Biomedical Language Processing (BioNLP) · doi:10.18653/v1/2024.bionlp-1.34 · Cited by 3
Ling, X., Cai, C., Kong, D., Wei, Z., Wu, J., Wang, L., & Deng, Z. (2025) · Journal of Chemical Information and Modeling · doi:10.1021/acs.jcim.5c00378 · Cited by 2
Secondary Structure-Guided Novel Protein Sequence Generation with Latent Graph Diffusion
Hu, Y., Tan, Y., Han, A., Zheng, L., Hong, L., & Zhong, B. (2024) · IEEE International Conference on Bioinformatics and Biomedicine (BIBM) · doi:10.1109/bibm62325.2024.10822117 · Cited by 4
TMvisDB: resource for transmembrane protein annotation and 3D visualizationPreprint
Marquet, C., Grekova, A., Houri, L., Bernhofer, M., Soto, L. F. J., Karl, T., Heinzinger, M., Dallago, C., & Rost, B. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.11.30.518551 · Cited by 2
Wang, Y., Wang, B., Shi, T., Fu, J., Zhou, Y., & Zhang, Z. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.11.06.565922 · Cited by 2
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Tan, Y., Zhong, B., Zheng, L., Fan, G., & Hong, L. (2024) · eLife · doi:10.7554/elife.98033.1 · Cited by 2
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Страшнов, П. В., Shevtsov, A., Meshchaninov, V., Ivanova, M., Nikolaev, F., Kardymon, O. L., & Vetrov, D. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.10.25.620213 · Cited by 3
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Pourmirzaei, M., Han, Y., Esmaili, F., Pourmirzaei, M., Alqarghuli, S., Chen, K., & Xu, D. H. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.03.03.641065 · Cited by 2
Nainani, J., Reimer, B. M., Watts, C., Jensen, D., & Green, A. G. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.08.22.671739 · Cited by 2
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Murphy, A., Kocherovsky, M., Dayan, N. A., Miralavy, I., Gilad, A. A., & Banzhaf, W. (2025) · Lecture notes in computer science · doi:10.1007/978-3-031-90065-5_30 · Cited by 1
Wang, L., Qiu, C., Xiao, L., & Chen, J. (2025) · IEEE International Conference on Bioinformatics and Biomedicine (BIBM) · doi:10.1109/bibm66473.2025.11356223 · Cited by 1
Wu, J., Wang, C., Zeng, Y., Song, Y.-Y., Xu, S., Nie, J., Wang, T.-X., Ma, J.-C., Feng, J., & Wu, L.-H. (2026) · Cell Reports · doi:10.1016/j.celrep.2026.117275 · Cited by 3
Das, A., Sen, S., Ahmadi, Z., & Debnath, M. (2026) · Chemical Engineering Journal · doi:10.1016/j.cej.2026.178296 · Cited by 1
Li, K., Ding, S., Guo, Z., Fan, Y., Liu, H., Sun, Y., Zhang, G., Zhang, R., Huang, L., & Zhou, F. (2025) · Expert Systems with Applications · doi:10.1016/j.eswa.2025.127786 · Cited by 1
Yu, Y., Jiang, F., Ma, X., Zhang, L., Zhong, B., Ouyang, W., Fan, G., Yu, H., Hong, L., & Li, M. (2025) · neural information processing systems · doi:10.52202/085713-4581 · Cited by 1
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Pokharel, S., Pratyush, P., Chaudhari, M., Heinzinger, M., Caragea, D., Saigo, H., & KC, D. B. (2025) · Methods in molecular biology · doi:10.1007/978-1-0716-4623-6_1 · Cited by 1
Gou, W., Tan, Y., Liu, C., Li, M., Fan, G., & Yu, H. (2025) · Journal of Chemical Information and Modeling · doi:10.1021/acs.jcim.5c00856 · Cited by 1
Hussain, M., Ou, Y., & Ho, Q. (2025) · Computational Biology and Chemistry · doi:10.1016/j.compbiolchem.2025.108595 · Cited by 1
Umar, M., Shamim, L., Musharaf, I., Tabasum, P., Malhotra, V., Farhan, K., Hidayat, A., Waqas, M., Anwar, A., Qadri, M., et al. (2025) · Journal of intelligent medicine. · doi:10.1002/jim4.70018 · Cited by 1
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Xia, Y., Sun, J., Du, X., Liang, Z., Wu, X., Shi, W., Shao, B., Guo, S., & Huo, Y. (2026) · Nucleic Acids Research · doi:10.1093/nar/gkag122 · Cited by 2
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Liang, J., Luo, Y., Su, B., & Peng, Z. (2026) · Briefings in Bioinformatics · doi:10.1093/bib/bbag126 · Cited by 1
Lin, Y., Wu, J., Yang, S., & Wang, Y. (2026) · Communications Chemistry · doi:10.1038/s42004-026-02062-y · Cited by 1
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Balbi, L., Sousa, R. T., Cotovio, P. G., & Pesquita, C. (2026) · Briefings in Bioinformatics · doi:10.1093/bib/bbag313 · Cited by 1
Huang, Y., Ge, R., Wu, L., Sun, Y., Wang, Z., & Cao, M. (2026) · ACS Synthetic Biology · doi:10.1021/acssynbio.6c00199 · Cited by 1
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Cui, T., Li, B., Cai, B., Wang, H., & Li, W. (2026) · Trends in Molecular Medicine · doi:10.1016/j.molmed.2026.06.004 · Cited by 1
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Min, G., & Peng, H. (2026) · Methods in molecular biology · doi:10.1007/978-1-0716-5249-7_13 · Cited by 1
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Jiao, X., Lin, S. Q., Fan, J., Liang, Z., Mao, W., Chen, H., & Shen, C. (2026) · Bioinformatics · doi:10.1093/bioinformatics/btag525 · Cited by 1
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Şahin, İ. O., Gökçe, N., Yildirim, D. T., Yildirim, A. B., Akalin, H., Martin, D. K., Beccari, T., Vicente, O., Radecka, I., Tchuenbou-Magaia, F. L., et al. (2026) · The EuroBiotech Journal · doi:10.2478/ebtj-2026-0015 · Cited by 1
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Xiong, J., Gaur, I., Lukarska, M., Nisonoff, H. M., Oltrogge, L. M., Savage, D. F., & Listgarten, J. (2026) · Nature Biotechnology · doi:10.1038/s41587-026-03207-z · Cited by 3
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Harding-Larsen, D., Lax, B. M., Weingarten, C. K., Sako, A., Mazurenko, S., & Welner, D. H. (2026) · Chem & Bio Engineering · doi:10.1021/cbe.6c00028 · Cited by 1
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Lopez, I. (. M. (2003) · doi:10.15581/10171/67243 · Cited by 1
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Prakash, M., Moskalev, A., DiMaggio, P. A., Combs, S., Mansi, T., Scheer, J. K., & Liao, R. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.10.15.618385
Protein Family Sequence Generation through ProGen2 Fine-Tuning
Hrbáň, H., & Hoksza, D. (2024) · IEEE International Conference on Bioinformatics and Biomedicine (BIBM) · doi:10.1109/bibm62325.2024.10822651
The Influence of Gendered Language on Leadership Decision-Making Styles
Al-Badawi, M. A. (2025) · Studies in systems, decision and control · doi:10.1007/978-3-031-76011-2_109
Semantical and Geometrical Protein Encoding Toward Enhanced Bioactivity and ThermostabilityPreprint
Tan, Y., Zhong, B., Zheng, L., Fan, G., & Hong, L. (2025) · eLife · doi:10.7554/elife.98033.3 · Cited by 1
CHIEF: An Attention-based Ensemble Learning Framework for Functional Protein DesignPreprint
Geng, Z., Wang, Y., Liu, T., Tan, A., Wu, S., Guo, X., Li, R., Hou, X., Sun, K., Wu, L., et al. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.03.07.641005
Protein Family Sequence Generation Through ProGen2 Fine-Tuning
Hrbáň, H., & Hoksza, D. (2024) · IEEE International Conference on Bioinformatics and Biomedicine (BIBM) · doi:10.1109/bibm62325.2024.10947607
PLM-OMG: Protein Language Model-Based Ortholog Detection for Cross-Species Cell Type MappingPreprint
Chau, T. N., & Li, S. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.05.30.657127
Varghese, D. M., Athulya, T., Mohani, V. K., & Ahmad, S. (2025) · Methods in molecular biology · doi:10.1007/978-1-0716-4623-6_13
Ranking Protein–Protein Models with Large Language Models and Graph Neural Networks
Xu, X., & Bonvin, A. M. J. J. (2025) · Methods in molecular biology · doi:10.1007/978-1-0716-4623-6_4
Metadata
- Authors
- Nijkamp, E., Ruffolo, J. A., Weinstein, E. N., Naik, N., & Madani, A.
- Affiliations
- Salesforce (United States) (US)
- Johns Hopkins University (US)
- Columbia University (US)
- Corresponding author
- Madani, Ali
- Venue
- Cell Systems
- Publisher
- Elsevier BV
- Published
- 2023-11
- Volume
- 14
- Issue
- 11
- Pages
- 968-978.e3
- ISSN
- 2405-4712 (ISSN-L)
- OpenAlex
- W4388024559
- Cited by
- 480
- References
- 57
- Citations by year
- 2026: 150 · 2025: 169 · 2024: 112 · 2023: 39
- FWCI
- 42.06
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- www.elsevier.com/tdm/userlicense/1.0/ (tdm)
- www.elsevier.com/legal/tdmrep-license (tdm)
- www.elsevier.com/open-access/userlicense/1.0/ (vor, 380 d)
- doi.org/10.15223/policy-017 (stm-asf)
- doi.org/10.15223/policy-037 (stm-asf)
- doi.org/10.15223/policy-012 (stm-asf)
- doi.org/10.15223/policy-029 (stm-asf)
- doi.org/10.15223/policy-004 (stm-asf)
- Locations
- Cell Systems (publishedVersion)
- PubMed (publishedVersion)
- arXiv (Cornell University) (submittedVersion)
- arXiv (Cornell University) (cc-by)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Machine Learning in Bioinformatics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Protein Structure and Dynamics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › RNA and protein synthesis mechanisms
- Keywords
- protein language models, protein sequence generation, protein fitness prediction, protein design, model scaling, data distribution, immune repertoire, attention-based models
- MeSH terms
- Amino Acid Sequence, Artificial Intelligence, Language, Proteins, Databases, Factual
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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