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Modeling the language of life – Deep Learning Protein Sequences
Open accessPreprint — published as 10.1186/s12859-019-3220-8 · Paper page
Abstract
Background: One common task in Computational Biology is the prediction of aspects of protein function and structure from their amino acid sequence. For 26 years, most state-of-the-art approaches toward this end have been marrying machine learning and evolutionary information. The retrieval of related proteins from ever growing sequence databases is becoming so time-consuming that the analysis of entire proteomes becomes challenging. On top, evolutionary information is less powerful for small families, e.g. for proteins from the Dark Proteome . Results: We introduce a novel way to represent protein sequences as continuous vectors ( embeddings ) by using the deep bi-directional model ELMo taken from natural language processing (NLP). The model has effectively captured the biophysical properties of protein sequences from unlabeled big data (UniRef50). After training, this knowledge is transferred to single protein sequences by predicting relevant sequence features. We refer to these new embeddings as SeqVec ( Seq uence-to- Vec tor) and demonstrate their effectiveness by training simple convolutional neural networks on existing data sets for two completely different prediction tasks. At the per-residue level, we significantly improved secondary structure (for NetSurfP-2.0 data set: Q3=79%±1, Q8=68%±1) and disorder predictions (MCC=0.59±0.03) over methods not using evolutionary information. At the per-protein level, we predicted subcellular localization in ten classes (for DeepLoc data set: Q10=68%±1) and distinguished membrane-bound from water-soluble proteins (Q2= 87%±1). All results built upon the embeddings gained from the new tool SeqVec neither explicitly nor implicitly using evolutionary information. Nevertheless, it improved over some methods using such information. Where the lightning-fast HHblits needed on average about two minutes to generate the evolutionary information for a target protein, SeqVec created the vector representation on average in 0.03 seconds. Conclusion: We have shown that transfer learning can be used to capture biochemical or biophysical properties of protein sequences from large unlabeled sequence databases. The effectiveness of the proposed approach was showcased for different prediction tasks using only single protein sequences. SeqVec embeddings enable predictions that outperform even some methods using evolutionary information. Thus, they prove to condense the underlying principles of protein sequences. This might be the first step towards competitive predictions based only on single protein sequences. Availability: SeqVec: https://github.com/mheinzinger/SeqVec Prediction server: https://embed.protein.properties
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Cites
Hochreiter, S., & Schmidhuber, J. (1997) · Neural Computation · doi:10.1162/neco.1997.9.8.1735 · Cited by 102,318
Adam: A Method for Stochastic Optimization
Kingma, D. P., & Ba, J. (2014) · arXiv · doi:10.48550/arxiv.1412.6980 · Cited by 98,946
Gapped BLAST and PSI-BLAST: a new generation of protein database search programs
Altschul, S. (1997) · Nucleic Acids Research · doi:10.1093/nar/25.17.3389 · Cited by 73,302
BERT: Pre-training of Deep Bidirectional Transformers for Language Understanding
Devlin, J., Chang, M.-W., Lee, K., & Toutanova, K. (2018) · arXiv · doi:10.48550/arxiv.1810.04805 · Cited by 73,173
Berman, H. M. (2000) · Nucleic Acids Research · doi:10.1093/nar/28.1.235 · Cited by 40,196
Glove: Global Vectors for Word Representation
Pennington, J., Socher, R., & Manning, C. (2014) · Proceedings of the 2014 Conference on Empirical Methods in Natural Language Processing (EMNLP) · doi:10.3115/v1/d14-1162 · Cited by 34,071
(2014) · Cited by 33,419
(2008) · Journal of Machine Learning Research · Cited by 33,051
Batch Normalization: Accelerating Deep Network Training by Reducing Internal Covariate Shift
Ioffe, S., & Szegedy, C. (2015) · arXiv · doi:10.48550/arxiv.1502.03167 · Cited by 27,818
Efficient Estimation of Word Representations in Vector Space
Mikolov, T., Chen, K., Corrado, G., & Dean, J. (2013) · arXiv · doi:10.48550/arxiv.1301.3781 · Cited by 26,568
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Items 201–300 of 599
Deep embeddings to comprehend and visualize microbiome protein space
Odrzywolek, K., Karwowska, Z., Majta, J., Byrski, A., Milanowska-Zabel, K., & Kosciolek, T. (2022) · Scientific Reports · doi:10.1038/s41598-022-14055-7 · Cited by 22
Pathogen genomic surveillance and the AI revolution
Lytras, S., Lamb, K. D., Ito, J., Grove, J., Yuan, K., Sato, K., Hughes, J., & Robertson, D. L. (2025) · Journal of Virology · doi:10.1128/jvi.01601-24 · Cited by 24
pLM4CPPs: Protein Language Model-Based Predictor for Cell Penetrating Peptides
Kumar, N., Du, Z., & Li, Y. (2025) · Journal of Chemical Information and Modeling · doi:10.1021/acs.jcim.4c01338 · Cited by 22
Large-scale predicting protein functions through heterogeneous feature fusion
Zheng, R., Huang, Z., & Deng, L. (2023) · Briefings in Bioinformatics · doi:10.1093/bib/bbad243 · Cited by 23
CollaPPI: A Collaborative Learning Framework for Predicting Protein-Protein Interactions
Ma, W., Bi, X., Jiang, H., Zhang, S., & Wei, Z. (2024) · IEEE Journal of Biomedical and Health Informatics · doi:10.1109/jbhi.2024.3375621 · Cited by 22
Filipavicius, M., Manica, M., Cadow, J., & Martínez, M. R. (2020) · arXiv (Cornell University) · doi:10.48550/arxiv.2012.03084 · Cited by 25
Fast and accurate protein intrinsic disorder prediction by using a pretrained language model
Song, Y., Yuan, Q., Chen, S., Chen, K., Zhou, Y., & Yang, Y. (2023) · Briefings in Bioinformatics · doi:10.1093/bib/bbad173 · Cited by 21
Wang, J., Quan, L., Jin, Z. X., Wu, H., Ma, X., Wang, X., Xie, J., Pan, D., Chen, T., Wu, T., et al. (2024) · IEEE Journal of Biomedical and Health Informatics · doi:10.1109/jbhi.2024.3400521 · Cited by 22
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Yang, W. P., Liu, Y., & Xiao, C. (2022) · Knowledge-Based Systems · doi:10.1016/j.knosys.2022.108356 · Cited by 23
Recent advances in generative biology for biotherapeutic discovery
Mock, M. L., LANGMEAD, C. J., Grandsard, P. J., Edavettal, S. C., & Russell, A. J. (2024) · Trends in Pharmacological Sciences · doi:10.1016/j.tips.2024.01.003 · Cited by 20
Assessing the role of evolutionary information for enhancing protein language model embeddings
Erckert, K., & Rost, B. (2024) · Scientific Reports · doi:10.1038/s41598-024-71783-8 · Cited by 21
Self-citationPhage therapy against Klebsiella pneumoniae: An evolving perspective
Xing, J., Zhang, H., Zheng, L., Zhao, J., Zhang, Y., Xu, Z., Zhai, Y., Hu, G., Li, J., & Wu, H. (2025) · Biotechnology Advances · doi:10.1016/j.biotechadv.2025.108689 · Cited by 22
Manfredi, M., Savojardo, C., Martelli, P. L., & Casadio, R. (2023) · Journal of Molecular Biology · doi:10.1016/j.jmb.2023.167963 · Cited by 20
Elucidating the functional roles of prokaryotic proteins using big data and artificial intelligence
Ardern, Z., Chakraborty, S., Lenk, F., & Kaster, A. (2023) · FEMS Microbiology Reviews · doi:10.1093/femsre/fuad003 · Cited by 22
HyperPCM: Robust Task-Conditioned Modeling of Drug–Target Interactions
Svensson, E., Hoedt, P.-J., Hochreiter, S., & Klambauer, G. (2024) · Journal of Chemical Information and Modeling · doi:10.1021/acs.jcim.3c01417 · Cited by 21
Hu, Y., Wang, Y., Hu, X., Chao, H., Li, S., Ni, Q., Zhu, Y., Hu, Y., Zhao, Z., & Chen, M. (2024) · Computational and Structural Biotechnology Journal · doi:10.1016/j.csbj.2024.01.015 · Cited by 23
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Bojar, D., & Fussenegger, M. (2019) · Small · doi:10.1002/smll.201903093 · Cited by 20
Mul-SNO: A Novel Prediction Tool for S-Nitrosylation Sites Based on Deep Learning Methods
Zhao, Q., Ma, J., Wang, Y., Xie, F., Lv, Z., Xu, Y., Shi, H., & Han, K. (2021) · IEEE Journal of Biomedical and Health Informatics · doi:10.1109/jbhi.2021.3123503 · Cited by 20
Self-citationMehdiabadi, M., Conte, A. D., Nugnes, M. V., Aspromonte, M. C., Tosatto, S. C. E., & Piovesan, D. (2025) · Proteins Structure Function and Bioinformatics · doi:10.1002/prot.70045 · Cited by 20
Villalobos-Alva, J., Ochoa-Toledo, L., Villalobos-Alva, M. J., Aliseda, A., Escamirosa, F. P., Altamirano-Bustamante, N. F., Ochoa-Fernández, F., Zamora-Solís, R., Villalobos-Alva, S., Revilla, M., et al. (2022) · Frontiers in Bioengineering and Biotechnology · doi:10.3389/fbioe.2022.788300 · Cited by 19
Ten quick tips for sequence-based prediction of protein properties using machine learning
Hou, Q., Waury, K., Gogishvili, D., & Feenstra, K. A. (2022) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1010669 · Cited by 20
When Protein Structure Embedding Meets Large Language Models
Ali, S., Chourasia, P., & Patterson, M. (2023) · Genes · doi:10.3390/genes15010025 · Cited by 15
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Merino, G. A., Saidi, R., Milone, D. H., Stegmayer, G. S., & Martin, M. J. (2022) · Bioinformatics · doi:10.1093/bioinformatics/btac536 · Cited by 19
LambdaPP : Fast and accessible protein‐specific phenotype predictions
(2022) · doi:10.1002/pro.4524 · Cited by 19
Self-citationDeep Neural Network Framework Based on Word Embedding for Protein Glutarylation Sites Prediction
Liu, C.-M., Ta, V.-D., Le, N. Q. K., Addis, D., & Shi, C. (2022) · Life · doi:10.3390/life12081213 · Cited by 18
TPepPro: a deep learning model for predicting peptide–protein interactions
Xiao-hong, J., Chen, Z., Yu, D., Jiang, Q., Chen, Z., Yan, B., Qin, J., Liu, Y., & Wang, J. (2024) · Bioinformatics · doi:10.1093/bioinformatics/btae708 · Cited by 18
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Armenteros, J. J. A., Johansen, A. R., Winther, O., & Nielsen, H. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.03.09.983585 · Cited by 19
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Hamelin, D. J., Scicluna, M., Saadie, I., Mostefai, F., Grenier, J., Baron, C., Caron, É., & Hussin, J. (2025) · Computational and Structural Biotechnology Journal · doi:10.1016/j.csbj.2025.03.044 · Cited by 21
Li, J., Tian, Y., Li, R., Zhou, T., Li, J., Ding, K., & Li, J. (2021) · Artificial Intelligence in Medicine · doi:10.1016/j.artmed.2021.102024 · Cited by 18
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Behjati, A., Zare‐Mirakabad, F., Arab, S. S., & Nowzari-Dalini, A. (2022) · Computational Biology and Chemistry · doi:10.1016/j.compbiolchem.2022.107717 · Cited by 22
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Murad, T., Ali, S., & Patterson, M. (2023) · Biology · doi:10.3390/biology12060854 · Cited by 17
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Wang, L., Zhang, H., Xu, W. M., Xue, Z., & Wang, Y. (2023) · Cell Reports Physical Science · doi:10.1016/j.xcrp.2023.101600 · Cited by 20
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Lyu, D., Wang, X., Chen, Y., & Wang, F. (2024) · iScience · doi:10.1016/j.isci.2024.109334 · Cited by 17
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Zhao, L., Li, J., Zhan, W., Jiang, X., & Zhang, B. (2024) · Scientific Reports · doi:10.1038/s41598-024-67403-0 · Cited by 17
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Hou, Z., Yang, Y., Li, H., Wong, K., & Li, X. (2021) · Briefings in Bioinformatics · doi:10.1093/bib/bbab288 · Cited by 17
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Elnaggar, A., Heinzinger, M., Dallago, C., & Rost, B. (2019) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/864405 · Cited by 17
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Morton, J. T., Strauss, C. E. M., Blackwell, R., Berenberg, D., Gligorijević, V., & Bonneau, R. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.11.03.365932 · Cited by 17
TM-Vec: template modeling vectors for fast homology detection and alignmentPreprint
Hamamsy, T., Morton, J. T., Berenberg, D., Carriero, N. J., Gligorijević, V., Blackwell, R., Strauss, C. E. M., Leman, J. K., Cho, K., & Bonneau, R. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.07.25.501437 · Cited by 16
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MMSMAPlus: a multi-view multi-scale multi-attention embedding model for protein function prediction
Wang, Z., Deng, Z., Zhang, W., Lou, Q., Choi, K., Wei, Z., Wang, L., & Wu, J. (2023) · Briefings in Bioinformatics · doi:10.1093/bib/bbad201 · Cited by 16
A novel hybrid CNN and BiGRU-Attention based deep learning model for protein function prediction
Sharma, L., Deepak, A., Ranjan, A., & Krishnasamy, G. (2023) · Statistical Applications in Genetics and Molecular Biology · doi:10.1515/sagmb-2022-0057 · Cited by 18
Application of Sequence Embedding in Protein Sequence-Based Predictions
Ibtehaz, N., & Kihara, D. (2022) · WORLD SCIENTIFIC eBooks · doi:10.1142/9789811258589_0002 · Cited by 18
Prediction of Protein–Protein Interaction Sites Based on Stratified Attentional Mechanisms
Tang, M., Wu, L., Yu, X., Chu, Z., Jin, S., & Liu, J. (2021) · Frontiers in Genetics · doi:10.3389/fgene.2021.784863 · Cited by 16
Mardikoraem, M., & Woldring, D. (2023) · Pharmaceutics · doi:10.3390/pharmaceutics15051337 · Cited by 16
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Breimann, S., Kamp, F., Steiner, H., & Frishman, D. (2024) · Journal of Molecular Biology · doi:10.1016/j.jmb.2024.168717 · Cited by 15
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Zhao, Y., Yang, Z., Hong, Y., Yang, Y., Wang, L., Zhang, Y., Lin, H., & Wang, J. (2023) · IEEE Transactions on NanoBioscience · doi:10.1109/tnb.2023.3278033 · Cited by 15
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Wang, Q., Wang, B., Xu, Z., Wu, J., Zhao, P., Li, Z., Wang, S., Huang, J., & Cui, S. R. (2021) · Proceedings of the AAAI Conference on Artificial Intelligence · doi:10.1609/aaai.v35i1.16141 · Cited by 18
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Tubiana, J., Schneidman‐Duhovny, D., & Wolfson, H. J. (2022) · Journal of Molecular Biology · doi:10.1016/j.jmb.2022.167758 · Cited by 15
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Self-citationBhushan, V., & Nita‐Lazar, A. (2024) · Journal of Proteome Research · doi:10.1021/acs.jproteome.3c00839 · Cited by 15
Yang, S., & Xu, P. (2024) · Analytical Biochemistry · doi:10.1016/j.ab.2024.115523 · Cited by 15
Kabir, A., Moldwin, A., Bromberg, Y., & Shehu, A. (2024) · Bioinformatics Advances · doi:10.1093/bioadv/vbae119 · Cited by 13
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Frolova, D., Pak, M., Litvin, A., Sharov, I., Ivankov, D., & Oseledets, I. (2024) · Bioinformatics Advances · doi:10.1093/bioadv/vbaf117 · Cited by 12
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Furui, K., Sakano, K., & Ohue, M. (2025) · Allergology International · doi:10.1016/j.alit.2025.08.004 · Cited by 12
NLP-BCH-Ens: NLP-based intelligent computational model for discrimination of malaria parasite
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SPRoBERTa: protein embedding learning with local fragment modeling
Wu, L., Yin, C., Zhu, J., Wu, Z., He, L., Xia, Y., Xie, S., Qin, T., & Liu, T. (2022) · Briefings in Bioinformatics · doi:10.1093/bib/bbac401 · Cited by 12
DeepBSRPred: deep learning-based binding site residue prediction for proteins
Nikam, R., Yugandhar, K., & Gromiha, M. M. (2022) · Amino Acids · doi:10.1007/s00726-022-03228-3 · Cited by 12
Lim, P. K., Julca, I., & Mutwil, M. (2023) · Computational and Structural Biotechnology Journal · doi:10.1016/j.csbj.2023.01.013 · Cited by 12
AI4Protein: transforming the future of protein design
Wang, D., Tan, Z., Gao, J., Zhang, S., Shen, J., & Lu, Y. (2025) · Science China Life Sciences · doi:10.1007/s11427-024-2906-3 · Cited by 12
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Lee, T.-H., Charchar, P., Sani, M., Le, D.-H., Le, T. C., Yarovsky, I., Separovic, F., & Aguilar, M. (2026) · Chemical Reviews · doi:10.1021/acs.chemrev.5c00994 · Cited by 12
MPI-VGAE: protein–metabolite enzymatic reaction link learning by variational graph autoencoders
Wang, C., Yuan, C., Wang, Y., Chen, R., Shi, Y., Zhang, T., Xue, F., Patti, G. J., Wei, L., & Hou, Q. (2023) · Briefings in Bioinformatics · doi:10.1093/bib/bbad189 · Cited by 13
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Paul, D., Saha, S., Basu, S., & Chakraborti, T. (2024) · Scientific Reports · doi:10.1038/s41598-024-69617-8 · Cited by 11
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Maier, A. J., Cha, M., Burgess, S., Wang, A., Cuellar, C., Kim, S. Y., Rajan, N. S., Neyyan, J., SENGUPTA, R., O’Connor, K., et al. (2025) · mAbs · doi:10.1080/19420862.2024.2439988 · Cited by 11
Allman, B. E., Vieira, L., Diaz, D. J., & Wilke, C. O. (2025) · Journal of The Royal Society Interface · doi:10.1098/rsif.2024.0598 · Cited by 11
Asim, M. N., Asif, T., Hassan, F., & Dengel, A. (2025) · Database · doi:10.1093/database/baaf027 · Cited by 11
Xu, H., Hu, R., & Zhao, Z. (2023) · Molecular Biology and Evolution · doi:10.1093/molbev/msad253 · Cited by 11
Cuturello, F., Celoria, M., Ansuini, A., & Cazzaniga, A. (2024) · Bioinformatics · doi:10.1093/bioinformatics/btae447 · Cited by 11
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Locality-aware pooling enhances protein language model performance across varied applications
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Georgoulis, E., Zervou, M. A., & Pantazis, Y. (2025) · Scientific Reports · doi:10.1038/s41598-025-21223-y · Cited by 13
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PROVAL: A framework for comparison of protein sequence embeddings
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Metadata
- Authors
- Heinzinger, M., Elnaggar, A., Wang, Y., Dallago, C., Nechaev, D., Matthes, F., & Rost, B.
- Affiliations
- Technical University of Munich (DE)
- Leibniz Supercomputing Centre (DE)
- Corresponding author
- Heinzinger, Michael
- Venue
- BMC Bioinformatics
- Publisher
- openRxiv
- Published
- 2019-04-19
- Volume
- 20
- Issue
- 1
- Pages
- 723-723
- ISSN
- 1471-2105 (ISSN-L)
- PMC
- PMC6918593
- OpenAlex
- W2995514860
- Cited by
- 606
- References
- 105
- Citations by year
- 2026: 53 · 2025: 103 · 2024: 115 · 2023: 117 · 2022: 99 · 2021: 83 · 2020: 29 · 2019: 7
- FWCI
- 18.03
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- cc-by
- Locations
- BMC Bioinformatics (publishedVersion, cc-by)
- bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by-nc-nd)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- mediaTUM – the media and publications repository of the Technical University Munich (Technical University Munich) (submittedVersion, other-oa)
- Columbia Academic Commons (Columbia University)
- Funders
- Alexander von Humboldt-Stiftung
- Nvidia
- Deutsche Forschungsgemeinschaft (DFG-GZ: RO1320/4-1, RO1320/4‐1)
- Bundesministerium für Bildung und Forschung
- Leibniz-Gemeinschaft
- 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 › Fractal and DNA sequence analysis
- Keywords
- protein sequence embeddings, ELMo, transfer learning, protein function prediction, protein structure prediction, secondary structure prediction, subcellular localization prediction, membrane proteins, dark proteome, metaproteomics, microbiome analysis
- MeSH terms
- Machine Learning, Amino Acid Sequence, Natural Language Processing, Proteins, Neural Networks, Computer, Sequence Analysis, Computational Biology, Databases, Nucleic Acid, Databases, Protein, Proteomics
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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