The DOI matches its Crossref record; 177 of the 177 references it cites were found in a registry or an index; 65 of the 86 sources its claims cite were checked for support.
Sequence modeling and design from molecular to genome scale with Evo
Open accessPublished version of a preprint
Abstract
The genome is a sequence that encodes the DNA, RNA, and proteins that orchestrate an organism’s function. We present Evo, a long-context genomic foundation model with a frontier architecture trained on millions of prokaryotic and phage genomes, and report scaling laws on DNA to complement observations in language and vision. Evo generalizes across DNA, RNA, and proteins, enabling zero-shot function prediction competitive with domain-specific language models and the generation of functional CRISPR-Cas and transposon systems, representing the first examples of protein-RNA and protein-DNA codesign with a language model. Evo also learns how small mutations affect whole-organism fitness and generates megabase-scale sequences with plausible genomic architecture. These prediction and generation capabilities span molecular to genomic scales of complexity, advancing our understanding and control of biology.
Overview
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
62 Own result · 28 Own interpretation · 16 Partly supported · 11 Own method · 10 Supported · 5 Couldn’t confirm · 5 Sources not found · 4 No full text of the sources · 4 Not checked yet · 4 No basis given · 1 Supported so far · accuracy not yet measured
Every source the paper cites for this claim was checked and supports it.
In the paper
“Evo also reaches competitive performance with leading protein-specific language models (41, 46–48) (Fig. 2B).”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting mutational effects on protein function
What the sources say
- Supportedcites [41] Meier J., “ Language models enable zero-shot prediction of the effects of mutations on protein function” in Advances in Neural Information Processing Systems, vol. 34, Ranzato M, Beygelzimer A, Dauphin Y, Liang PS, Wortman Vaughan J, Eds. (Curran Associates, Inc., 2021), pp. 29287– 29303. source
The Introduction (p. 2) identifies this paper as introducing ESM-1v, a protein language model, and §4.2 (p. 6) compares it with other protein language models. The citation supports the identification of a protein-specific model used as a comparator; the statement about Evo’s competitive performance is the citing authors’ own result, attributed to their Fig. 2B. (LLM, conf=0.88)
- Supportedcites [46] Yang KK, Fusi N, Lu AX, Convolutions are competitive with transformers for protein sequence pretraining. Cell Syst. 15, 286– 294.e2 (2024). doi: 10.1016/j.cels.2024.01.008 38428432 source
The citation identifies a protein-specific language model used as a comparator, rather than attributing Evo’s measured performance to this paper. In “CARP achieves comparable performance on downstream tasks to ESM” (page unavailable), the paper presents CARP and reports that it performs comparably to other protein language models across diverse tasks. The Evo comparison is the citing authors’ result, indicated by their Fig. 2B. (LLM, conf=0.85)
- Supportedcites [47] Lin Z., Evolutionary-scale prediction of atomic-level protein structure with a language model. Science 379, 1123– 1130 (2023). doi: 10.1126/science.ade2574 36927031 source
Section 2 (p. 2) identifies ESM-2 as a protein language model and reports that it outperforms other recent protein language models on structure-prediction benchmarks. This supports citing the paper as an example of a leading protein-specific model used for comparison; the Evo performance result belongs to the citing paper’s Fig. 2B. (LLM, conf=0.90)
- Supportedcites [48] Nijkamp E, Ruffolo JA, Weinstein EN, Naik N, Madani A, ProGen2: Exploring the boundaries of protein language models. Cell Syst. 14, 968– 978.e3 (2023). doi: 10.1016/j.cels.2023.10.002 37909046 source
The citation identifies ProGen2 as an example of a leading protein-specific language model, rather than attributing Evo’s comparison to the ProGen2 paper. The Discussion describes ProGen2 as a protein language model competitive with state-of-the-art methods across varied protein sequence landscapes. The cited paper does not itself evaluate Evo. (LLM, conf=0.88)
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Supported
Every source the paper cites for this claim was checked and supports it.
In the paper
“On DMS datasets of prokaryotic proteins, Evo’s zero-shot performance exceeded all other nucleotide models tested (Fig. 2B and table S4), including GenSLM (15)”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting mutational effects on protein function
What the sources say
- Supportedcites [15] Zvyagin M., GenSLMs: Genome-scale language models reveal SARS-CoV-2 evolutionary dynamics. Int. J. High Perform. Comput. Appl 37, 683– 705 (2023). doi: 10.1177/10943420231201154 source
The citation is attached to “GenSLM” and identifies the model used as a comparator, rather than sourcing the Evo benchmark result. Section 8 (Implications) identifies GenSLM as a nucleotide-sequence language model; Section 5.2.1 describes its foundation models. The cited paper need not report Evo’s DMS results to support this identification. (LLM, conf=0.91)
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Supported
Every source the paper cites for this claim was checked and supports it.
In the paper
“We found that Evo again outperforms all other tested nucleotide language models at this task, including RNA-FM (50), an RNA language model that is explicitly trained on ncRNA sequences”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting mutational effects on ncRNA function
What the sources say
- Supportedcites [50] Chen J., Interpretable RNA foundation model from unannotated data for highly accurate RNA structure and function predictions. arXiv: 2204. 00300 [q-bio.QM] (2022). source
The citation identifies RNA-FM, rather than attributing the citing authors’ Evo comparison to this source. The Introduction (p. 2) describes RNA-FM as a model trained on 23 million ncRNA sequences using self-supervised learning, supporting its identification as an RNA language model trained on ncRNA. The source does not establish the Evo comparison, but the citing sentence presents that as its authors’ own finding. (LLM, conf=0.93)
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Supported
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“Recombinant expression and purification of EvoCas9–1 paired with chemically synthesized Evo-generated single guide RNA (sgRNA) exhibited comparable in vitro cleavage activity to SpCas9 paired with the canonical SpCas9 sgRNA”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
What the sources say
- Partly supportedcites [62] Jinek M., A programmable dual-RNA-guided DNA endonuclease in adaptive bacterial immunity. Science 337, 816– 821 (2012). doi: 10.1126/science.1225829 22745249 PMC6286148 sourceExperimental — accuracy not yet measured
The single-guide-RNA cleavage passage (labeled Abstract; page unavailable) reports that S. pyogenes Cas9 guided by a chimeric RNA cleaved DNA in vitro. It does not report EvoCas9–1, an Evo-generated sgRNA, or a comparison of their cleavage activity with SpCas9. (LLM, conf=0.87)
- Could not verifycites [63] Hsu PD, DNA targeting specificity of RNA-guided Cas9 nucleases. Nat. Biotechnol 31, 827– 832 (2013). doi: 10.1038/nbt.2647 23873081 PMC3969858 sourceExperimental — accuracy not yet measured
The paper discusses SpCas9 guide-RNA architecture and cleavage activity, and its Online Methods describe an in vitro SpCas9 cleavage assay (page unavailable). Our copy does not address EvoCas9-1 or an Evo-generated sgRNA, so it cannot verify the claimed comparison; searches for those terms and for chemically synthesized sgRNA found no matching passage. (LLM, conf=0.78)
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Partly supported
Every source the paper cites for this claim was checked and supports it.
In the paper
“Combining the Evo embeddings with a supervised CNN architecture (mean Spearman r = 0.56) also approached the performance of Promoter Calculator (52), a state-of-the-art method for promoter activity prediction (mean Spearman r = 0.62).”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting activity of regulatory DNA
What the sources say
- Supportedcites [52] LaFleur TL, Hossain A, Salis HM, Automated model-predictive design of synthetic promoters to control transcriptional profiles in bacteria. Nat. Commun 13, 5159 (2022). doi: 10.1038/s41467-022-32829-5 36056029 PMC9440211 source
The citation is attached to “Promoter Calculator,” identifying the method used as the comparator. The source’s Discussion describes the Promoter Calculator as its model for predicting transcription initiation rates. The citing paper’s Spearman correlation comparison is not attributed to this source by that citation. (LLM, conf=0.90)
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Supported
The cited sources that could be checked couldn’t confirm or dispute this claim.
In the paper
“The AlphaFold3 cofolded structure prediction for EvoCas9–1 has a high mean pLDDT score of 90 across its protein, RNA, and DNA components”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
What the sources say
- Could not verifycites [64] Abramson J., Accurate structure prediction of biomolecular interactions with AlphaFold 3. Nature 630, 493– 500 (2024). doi: 10.1038/s41586-024-07487-w 38718835 PMC11168924 sourceExperimental — accuracy not yet measured
The ‘Predicted confidences track accuracy’ section explains pLDDT and illustrates it with a prediction of PDB 7T82, not EvoCas9–1. Searches for EvoCas9–1 and a mean pLDDT of 90 across protein, RNA, and DNA found no passage reporting that result in our copy. (LLM, conf=0.73)
plainSingle case- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Could not verify
The cited sources that could be checked couldn’t confirm or dispute this claim.
In the paper
“We observed that the model uses information from one end to specify the other end across a distance of ~1 to 2 kb, reflecting the model’s understanding of the tight evolutionary linkage of the two terminal elements”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of transposon systems
What the sources say
- Could not verifycites [70] Zhang Z., Protein language models learn evolutionary statistics of interacting sequence motifs. Proc. Natl. Acad. Sci. U.S.A 121, e2406285121 (2024). doi: 10.1073/pnas.2406285121 PMC11551344 39467119 sourceExperimental — accuracy not yet measured
The Results section, “Language Models Predict Structures by Looking Up Segment Pairings,” examines ESM-2 recovery of contacts between protein segments, not Evo’s prediction of IS200/IS605 terminal elements. Searches for IS200/IS605, Evo, and terminal-element linkage found no passage addressing the claimed 1–2 kb relationship. (LLM, conf=0.76)
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Could not verify
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“We also use Evo to generate genome-scale sequences with plausible high-level architecture more than1 megabase (Mb) in length, a scale that is orders of magnitude greater than previous methods (10, 13, 14).”
§ Sequence modeling and design from molecular to genome scale with Evo > Abstract > CONCLUSION:
What the sources say
- Partly supportedcites [10] Watson JL, De novo design of protein structure and function with RFdiffusion. Nature 620, 1089– 1100 (2023). doi: 10.1038/s41586-023-06415-8 37433327 PMC10468394 sourceExperimental — accuracy not yet measured
The Discussion reports RFdiffusion protein designs up to 600 residues, supporting its use as an example of a previous method operating at a much smaller scale. The paper does not report Evo generating genome-scale sequences longer than 1 Mb or assess their high-level architecture; it studies protein design rather than genome generation. (LLM, conf=0.83)
- Supportedcites [13] DaSilva LF, DNA-Diffusion: Leveraging Generative Models for Controlling Chromatin Accessibility and Gene Expression via Synthetic Regulatory Elements. bioRxiv 2024.02.01.578352 [Preprint] (2024); 10.1101/2024.02.01.578352. source
The citation is offered as an example of a previous sequence-generation method, not as evidence for Evo’s own results. The Abstract states that DNA-Diffusion generates 200 bp regulatory elements, orders of magnitude shorter than the citing sentence’s more-than-1-Mb sequences. (LLM, conf=0.90)
- Supportedcites [14] Lal A, Garfield D, Biancalani T, Eraslan G, regLM: Designing realistic regulatory DNA with autoregressive language models. bioRxiv 2024.02.14.580373 [Preprint] (2024); 10.1101/2024.02.14.580373. PMC11529870 39322281 source
The citation supports the comparison with a previous method, not the citing authors’ report of what Evo generated. Section 2.2 describes regLM work on 80-bp yeast promoters, and Section 2.3 describes 200-bp human enhancers; a sequence longer than 1 Mb is orders of magnitude longer than either. Our copy gives no page numbers for these passages. (LLM, conf=0.90)
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Partly supported
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“Previous work has shown that improvement beyond this performance range is difficult for protein language models with self-supervised pretraining alone (49)”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting mutational effects on protein function
What the sources say
- Partly supportedcites [49] Li F-Z, Amini AP, Yue Y, Yang KK, Lu AX, Feature Reuse and Scaling: Understanding Transfer Learning with Protein Language Models. bioRxiv 2024.02.05.578959 [Preprint] (2024); 10.1101/2024.02.05.578959. sourceExperimental — accuracy not yet measured
The Results (section 5; page unavailable) say that performance on many downstream tasks will not improve as protein language models improve. The Discussion (section 6; page unavailable) similarly says that simply scaling existing pretraining strategies will not improve performance for many tasks. The paper does not establish a general ceiling for all tasks or identify the particular performance range referenced in the citing sentence. (LLM, conf=0.82)
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Partly supported
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“language models specifically trained on large corpuses of protein sequences or nucleotide coding sequences have demonstrated an ability to predict mutational effects on protein function (41–43) without any task-specific fine-tuning or supervision.”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting mutational effects on protein function
What the sources say
- Supportedcites [41] Meier J., “ Language models enable zero-shot prediction of the effects of mutations on protein function” in Advances in Neural Information Processing Systems, vol. 34, Ranzato M, Beygelzimer A, Dauphin Y, Liang PS, Wortman Vaughan J, Eds. (Curran Associates, Inc., 2021), pp. 29287– 29303. source
The Introduction (p. 2) states that language models trained on large, diverse protein-sequence databases predict experimental measurements of protein function without further supervision and perform zero-shot prediction of mutational effects. This supports the protein-sequence alternative in the citing sentence without requiring task-specific fine-tuning. (LLM, conf=0.92)
- Supportedcites [42] Notin P., “ Tranception: Protein Fitness Prediction with Autoregressive Transformers and Inference-time Retrieval” in Proceedings of the 39th International Conference on Machine Learning, vol. 162, Chaudhuri K. et al., Eds. (PMLR, 2022), pp. 16990– 17017. source
The Introduction (p. 1) describes Tranception as pretrained on large quantities of protein sequences and reports its mutation-fitness prediction performance. Section 3.4 (p. 4) says the model is trained self-supervised and scores mutated sequences against wild type, supporting prediction without task-specific supervision or fine-tuning. This source supports the protein-sequence example in the jointly cited sentence; it does not need to establish the nucleotide-coding-sequence example. (LLM, conf=0.90)
- Partly supportedcites [43] Benegas G, Albors C, Aw AJ, Ye C, Song YS, GPN-MSA: an alignment-based DNA language model for genome-wide variant effect prediction. bioRxiv 2023.10.10.561776 [Preprint] (2024); 10.1101/2023.10.10.561776. sourceExperimental — accuracy not yet measured
The Introduction (page unavailable) says protein language models learn from massive sequence databases and accurately score missense variants in an unsupervised manner. It does not establish that models trained specifically on nucleotide coding sequences predict effects on protein function without task-specific fine-tuning; its discussion of DNA models concerns genome-wide variant effects. (LLM, conf=0.84)
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Supported
- It may state two separate claims as one. 44
- Probably not a substantial claim. 29
- The quote only partly states this claim. 2
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“We compiled a large genome dataset called OpenGenome (Materials and methods) with more than 80,000 bacterial and archaeal genomes and millions of predicted phage and plasmid sequences, covering 300 billion nucleotide tokens”
§ Sequence modeling and design from molecular to genome scale with Evo > Training Evo at scale on OpenGenome
What the sources say
- Could not verifycites [35] Parks DH, GTDB: An ongoing census of bacterial and archaeal diversity through a phylogenetically consistent, rank normalized and complete genome-based taxonomy. Nucleic Acids Res. 50, D785– D794 (2022). doi: 10.1093/nar/gkab776 34520557 PMC8728215 sourceExperimental — accuracy not yet measured
The Introduction and Data Availability sections describe GTDB as a bacterial and archaeal genome taxonomy, not OpenGenome or Evo’s training data. Searches for OpenGenome, 80,000 genomes, phage and plasmid sequences, and 300 billion nucleotide tokens found no passage establishing the claimed dataset composition or size. (LLM, conf=0.76)
- Partly supportedcites [36] Camargo AP, IMG/VR v4: An expanded database of uncultivated virus genomes within a framework of extensive functional, taxonomic, and ecological metadata. Nucleic Acids Res. 51, D733– D743 (2023). doi: 10.1093/nar/gkac1037 36399502 PMC9825611 sourceExperimental — accuracy not yet measured
The 'Automatic virus identification from IMG datasets' section (p. 3) says the authors processed 83,858 bacterial and archaeal isolate genomes, and the Abstract (p. 2) reports more than 15 million viral genomes and fragments. The paper does not establish that those sequences formed OpenGenome or were used to train Evo, nor does it give a 300-billion-token total. The Conclusion (p. 11) describes a plasmid-focused database as still in development, rather than reporting millions of plasmid sequences in this collection. (LLM, conf=0.86)
- Not checked — no full text of the sourcecites [37] Camargo AP, IMG/PR: A database of plasmids from genomes and metagenomes with rich annotations and metadata. Nucleic Acids Res. 52, D164– D173 (2024). doi: 10.1093/nar/gkad964 37930866 PMC10767988 source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Could not verify
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“existing attempts to model DNA as a language (15–17) are limited by the prevailing dense Transformer architecture, which incurs high computational cost as input sequence lengths grow relative to model width (scaling quadratically) and generally underperforms at single-nucleotide or byte-level resolution compared with models trained at coarser resolutions (18).”
§ Sequence modeling and design from molecular to genome scale with Evo > Abstract > CONCLUSION:
What the sources say
- Partly supportedcites [15] Zvyagin M., GenSLMs: Genome-scale language models reveal SARS-CoV-2 evolutionary dynamics. Int. J. High Perform. Comput. Appl 37, 683– 705 (2023). doi: 10.1177/10943420231201154 sourceExperimental — accuracy not yet measured
Split into 2 atoms — 0 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 1 unverifiable. Expand the claim to see per-atom verdicts.
- Partly supportedcites [16] Dalla-Torre H., The Nucleotide Transformer: Building and Evaluating Robust Foundation Models for Human Genomics. bioRxiv 2023.01.11.523679 [Preprint] (2023); 10.1101/2023.01.11.523679. PMC11810778 39609566 sourceExperimental — accuracy not yet measured
Split into 2 atoms — 1 confirmed, 0 partially supported, 0 disputed, 0 misattributed, 1 unverifiable. Expand the claim to see per-atom verdicts.
- Partly supportedcites [17] Zhou Z, Ji Y, Li W, Dutta P, Davuluri R, Liu H, DNABERT-2: Efficient foundation model and benchmark for multi-species genome. arXiv:2306.15006 [q-bio.GN] (2023). sourceExperimental — accuracy not yet measured
Split into 2 atoms — 1 confirmed, 0 partially supported, 0 disputed, 0 misattributed, 1 unverifiable. Expand the claim to see per-atom verdicts.
- Partly supportedcites [18] Tay Y., Charformer: Fast Character Transformers via Gradient-based Subword Tokenization. arXiv: 2106. 12672 [cs.CL] (2022). sourceExperimental — accuracy not yet measured
Split into 2 atoms — 0 confirmed, 2 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Partly supported
The sources with a verdict support this claim; the rest weren’t checked yet, have no full text we could read, or their verdicts aren’t rated yet.
In the paper
“For a few genomes, the zero-shot performance with 66k context is notably strong, with an AUROC of 0.90 on lambda phage essentiality data (74) and an AUROC of 0.84 on Pseudomonas aeruginosa essentiality data (75)”
§ Sequence modeling and design from molecular to genome scale with Evo > Learning gene essentiality with long genomic context
What the sources say
- Supportedcites [74] Piya D., Systematic and scalable genome-wide essentiality mapping to identify nonessential genes in phages. PLOS Biol. 21, e3002416 (2023). doi: 10.1371/journal.pbio.3002416 PMC10695390 38048319 source
The citation identifies the lambda-phage essentiality dataset used for the citing authors’ zero-shot evaluation, not the source of the AUROC result. The Results section (‘Genome-wide CRISPRi to map gene essentiality in λ’) reports gene-level essentiality measurements for lambda phage. The Pseudomonas result has a separate citation. (LLM, conf=0.91)
- Not checked — no full text of the sourcecites [75] Turner KH, Wessel AK, Palmer GC, Murray JL, Whiteley M, Essential genome of Pseudomonas aeruginosa in cystic fibrosis sputum. Proc. Natl. Acad. Sci. U.S.A 112, 4110– 4115 (2015). doi: 10.1073/pnas.1419677112 25775563 PMC4386324 source
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Supported
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“Evo’s zero-shot correlation on promoter-RBS sequences is also higher than the GC content of the promoter-RBS sequences (Spearman r = 0.47), zero-shot GenSLM likelihoods (Spearman r = 0.11), and RBS Calculator (Spearman r = 0.39)”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting activity of regulatory DNA
What the sources say
- Supportedcites [57] Salis HM, Mirsky EA, Voigt CA, Automated design of synthetic ribosome binding sites to control protein expression. Nat. Biotechnol 27, 946– 950 (2009). doi: 10.1038/nbt.1568 19801975 PMC2782888 source
The citation identifies RBS Calculator as a comparison tool. The paper’s Materials and Methods section (‘Software Implementation’; page unavailable) explicitly names its software the RBS Calculator and says it predicts translation initiation rates. The citation does not need to establish the citing paper’s Evo benchmark correlations. (LLM, conf=0.90)
- Partly supportedcites [58] Reis AC, Salis HM, An automated model test system for systematic development and improvement of gene expression models. ACS Synth. Biol 9, 3145– 3156 (2020). doi: 10.1021/acssynbio.0c00394 33054181 sourceExperimental — accuracy not yet measured
The section “A systematic comparison of translation initiation rate models” (page unavailable) identifies RBS Calculator as a model for predicting translation initiation rates. It does not report a Spearman correlation of 0.39 on promoter-RBS sequences or compare the model with Evo, GC content, or GenSLM; those performance claims are not supported by this source. (LLM, conf=0.86)
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Supported
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“As a result, Transformer-based DNA models are constrained to short context lengths and use schemes that aggregate nucleotides into the basic units of language models, called tokens, thereby sacrificing single-nucleotide resolution (15, 16, 21–23).”
§ Sequence modeling and design from molecular to genome scale with Evo > Abstract > CONCLUSION:
What the sources say
- Partly supportedcites [15] Zvyagin M., GenSLMs: Genome-scale language models reveal SARS-CoV-2 evolutionary dynamics. Int. J. High Perform. Comput. Appl 37, 683– 705 (2023). doi: 10.1177/10943420231201154 sourceExperimental — accuracy not yet measured
Section 4.2 contrasts standard 1,024- or 2,048-token contexts with the longer sequences needed for genomes, and Section 5.2.1 says GenSLM groups three nucleotides into each codon-level input token. But the paper does not say this sacrifices the ability to model individual nucleotide changes; Section 5.2.1 says GenSLM models individual mutations at nucleotide scale. The claim therefore overstates the resolution consequence and treats a common context-length challenge as a constraint on Transformer-based DNA models generally. (LLM, conf=0.83)
- Supportedcites [16] Dalla-Torre H., The Nucleotide Transformer: Building and Evaluating Robust Foundation Models for Human Genomics. bioRxiv 2023.01.11.523679 [Preprint] (2023); 10.1101/2023.01.11.523679. PMC11810778 39609566 source
The Discussion (page unavailable) says the models’ attention span is limited to 6 kb and explains that standard transformer attention makes much longer inputs intractable. Appendix A.3 (page unavailable) says the tokenizer normally represents six nucleotides as one token, supporting the claim about aggregation rather than single-nucleotide token resolution. (LLM, conf=0.88)
- Supportedcites [21] Fishman V., GENA-LM: A family of open-source foundational DNA language models for long sequences. bioRxiv 2023.06.12.544594 [Preprint] (2024); 10.1101/2023.06.12.544594. PMC11734698 39817513 source
The Introduction says DNABERT’s 500-base-pair input is insufficient for applications requiring longer context. The Conclusions say even GENA-LM’s longer inputs remain insufficient for some tasks and that BPE tokenization limits prediction granularity to tokens. Section 3.1 describes tokens that aggregate multiple base pairs, supporting the single-nucleotide-resolution point. (LLM, conf=0.88)
- Partly supportedcites [22] Ji Y, Zhou Z, Liu H, Davuluri RV, DNABERT: Pre-trained Bidirectional Encoder Representations from Transformers model for DNA-language in genome. Bioinformatics 37, 2112– 2120 (2021). doi: 10.1093/bioinformatics/btab083 33538820 PMC11025658 sourceExperimental — accuracy not yet measured
Methods §2.2 says DNABERT uses k-mer tokens rather than individual bases and accepts at most 512 tokens per input, supporting the tokenization and context-limit portions of the claim. But the same section describes a method for processing longer sequences, and the Results state that DNABERT enables visualization of nucleotide-level importance. The paper therefore does not support the unqualified claim that token aggregation sacrifices single-nucleotide resolution. (LLM, conf=0.88)
- Partly supportedcites [23] Hwang Y, Cornman AL, Kellogg EH, Ovchinnikov S, Girguis PR, Genomic language model predicts protein co-regulation and function. Nat. Commun 15, 2880 (2024). doi: 10.1038/s41467-024-46947-9 38570504 PMC10991518 sourceExperimental — accuracy not yet measured
The Introduction (page unavailable) describes GenSLMs as using codon-level representations and says that learning gene-to-gene interactions requires longer genomic segments. That supports an example of nucleotide aggregation and a need for longer context, but it does not establish that Transformer-based DNA models generally have short context lengths or sacrifice single-nucleotide resolution. (LLM, conf=0.78)
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Partly supported
Every source the paper cites for this claim was checked and supports it.
In the paper
“The DNA-targeting Cas9 nuclease is typically encoded within 3000 to 4800 base pairs (bp) of coding sequence and found in close genomic proximity to its cognate CRISPR array (60).”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
What the sources say
- Supportedcites [60] Hsu PD, Lander ES, Zhang F, Development and applications of CRISPR-Cas9 for genome engineering. Cell 157, 1262– 1278 (2014). doi: 10.1016/j.cell.2014.05.010 24906146 PMC4343198 source
In “Metagenomic, Structural, and Functional Diversity of Cas9” (page unavailable), the paper reports Cas9 protein lengths of roughly 900–1,600 amino acids, with most centered around 1,100 or 1,350 amino acids. That is consistent with the approximate 3,000–4,800 bp coding range. “CRISPR-Cas9: From Yogurt to Genome Editing” (page unavailable) says cas genes are typically adjacent to CRISPR repeat elements, supporting the proximity claim. (LLM, conf=0.87)
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
- Supported
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“even a single-nucleotide mutation in an essential gene can be incompatible with life if it disrupts that gene’s expression or function (72).”
§ Sequence modeling and design from molecular to genome scale with Evo > Learning gene essentiality with long genomic context
What the sources say
- Partly supportedcites [72] Rocha EPC, Danchin A, Gene essentiality determines chromosome organisation in bacteria. Nucleic Acids Res. 31, 6570– 6577 (2003). doi: 10.1093/nar/gkg859 14602916 PMC275555 sourceExperimental — accuracy not yet measured
The Introduction describes essential genes as serving indispensable cellular functions and discusses identifying them through gene inactivation. The ‘Essentiality drives gene strand bias’ section says nonfunctional peptides are especially counter-selected in essential functions. Neither section addresses whether a single-nucleotide mutation disrupting an essential gene’s expression or function can be incompatible with life. (LLM, conf=0.82)
plainconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
- Partly supported
Every source the paper cites for this claim was checked and supports it.
In the paper
“Diverse families of CRISPR-Cas systems are found throughout bacterial and archaeal life, such as Cas12- or Cas13-based systems that target DNA and RNA, respectively (61).”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
What the sources say
- Supportedcites [61] Koonin EV, Makarova KS, Origins and evolution of CRISPR-Cas systems. Phil. Trans. R. Soc. B 374, 20180087 (2019). doi: 10.1098/rstb.2018.0087 PMC6452270 30905284 source
The Abstract identifies CRISPR-Cas as bacterial and archaeal adaptive immunity systems, and the Diversity and classification section describes their substantial diversity. The Class 2 effectors section identifies Cas12 as a type V effector in complexes with target DNA and states that Cas13 effectors cleave target RNA. Our copy provides no page numbers for these passages. (LLM, conf=0.92)
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
- Supported
The cited sources that could be checked couldn’t confirm or dispute this claim.
In the paper
“For example, directly sampling from a large natural language model typically produces sequences that are grammatically correct yet locally biased toward simpler sentence constructions and that are globally incoherent, especially at long lengths.”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
What the sources say
- Could not verifycites [83] Wei J., Finetuned language models are zero-shot learners. arXiv: 2109. 01652 [cs.CL] (2022). sourceExperimental — accuracy not yet measured
The Introduction (p. 2) discusses large language models’ zero-shot task performance, not the quality of text produced by direct sampling. Searches for grammaticality, simpler sentence constructions, and global incoherence over long spans found no passage in our copy addressing those claims. (LLM, conf=0.75)
- Could not verifycites [84] Ouyang L., Training language models to follow instructions with human feedback. arXiv: 2203. 02155 [cs.CL] (2022). sourceExperimental — accuracy not yet measured
The Introduction (page unavailable) discusses factual errors, biased text, and failure to follow instructions, but not the claimed pattern in directly sampled text. Searches for grammaticality, simpler sentence constructions, and long-span incoherence found no passage in our copy that addresses the claim. (LLM, conf=0.73)
- Could not verifycites [85] Rafailov R., “ Direct Preference Optimization: Your Language Model is Secretly a Reward Model” in Advances in Neural Information Processing Systems, vol. 36, Oh A. et al., Eds. (Curran Associates, Inc., 2023), pp. 53728– 53741. sourceExperimental — accuracy not yet measured
The Introduction discusses why language-model outputs may need steering, but does not describe the claimed grammaticality, preference for simpler sentences, or long-span incoherence. The Additional information section discusses meaningless responses from an unlikelihood-training baseline, not the typical results of directly sampling a large language model. Searches for the claim’s distinctive wording and for long-text incoherence found no supporting passage in our copy; page numbers were not available. (LLM, conf=0.76)
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Could not verify
Every source the paper cites for this claim was checked and supports it.
In the paper
“TGA and TAA stop codons appeared most frequently, whereas TAG was least common, consistent with previously observed patterns in prokaryotic genomes (Fig. 6H) (80).”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
What the sources say
- Supportedcites [80] Korkmaz G, Holm M, Wiens T, Sanyal S, Comprehensive analysis of stop codon usage in bacteria and its correlation with release factor abundance. J. Biol. Chem 289, 30334– 30342 (2014). doi: 10.1074/jbc.M114.606632 25217634 PMC4215218 source
The Results (p. 3) report that TAA or TGA is the most frequent stop codon across the bacterial phyla examined, depending on phylum and genomic GC content, while TAG is generally the least frequent. This supports the cited comparison to prior prokaryotic patterns; the source does not need to describe the Evo-generated sequences themselves. (LLM, conf=0.93)
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Supported
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“We also examined stop codon frequencies across reading frames, a conserved genomic feature in prokaryotes (79).”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
What the sources say
- Partly supportedcites [79] Xu L, Kuo J, Liu J-K, Wong T-Y, Bacterial phylogenetic tree construction based on genomic translation stop signals. Microb. Inform. Exp 2, 6 (2012). doi: 10.1186/2042-5783-2-6 22651236 PMC3466146 sourceExperimental — accuracy not yet measured
The Discussion reports that related bacterial species often have similar stop-signal ratios in the second and third reading frames, and the Conclusion says signals across all three frames are interrelated. This supports a conserved pattern among related bacteria, but the paper does not establish that stop-codon frequencies are conserved across prokaryotes generally. (LLM, conf=0.83)
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Partly supported
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“Promisingly, in these domains, algorithmic techniques have emerged to improve the quality of generations compared with sampling from the pretrained model alone (83–85).”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
What the sources say
- Partly supportedcites [83] Wei J., Finetuned language models are zero-shot learners. arXiv: 2109. 01652 [cs.CL] (2022). sourceExperimental — accuracy not yet measured
The Introduction (p. 2) describes instruction tuning a pretrained language model, and the Results (p. 4) report that it significantly improves performance over the untuned model on most evaluated NLP tasks, including some generative tasks. The paper does not establish improved quality of open-ended generations compared with sampling from the pretrained model alone, and it does not study image generation. (LLM, conf=0.82)
- Partly supportedcites [84] Ouyang L., Training language models to follow instructions with human feedback. arXiv: 2203. 02155 [cs.CL] (2022). sourceExperimental — accuracy not yet measured
Section 3.1 describes fine-tuning a pretrained language model using demonstrations, human comparisons, and reward optimization. Section 4.1 reports that labelers preferred the resulting InstructGPT outputs to GPT-3 outputs on the paper’s test prompts. This supports improved natural-language generation over the pretrained model, but the cited paper does not establish the image-generation part of the claim. (LLM, conf=0.87)
- Partly supportedcites [85] Rafailov R., “ Direct Preference Optimization: Your Language Model is Secretly a Reward Model” in Advances in Neural Information Processing Systems, vol. 36, Oh A. et al., Eds. (Curran Associates, Inc., 2023), pp. 53728– 53741. sourceExperimental — accuracy not yet measured
The Related Work section says fine-tuning language models with human preferences improves performance on several text-generation tasks, and Section 6.2 reports better summarization results for DPO than for other methods. This supports the claim for natural-language generation, but the paper does not establish it for image generation. (LLM, conf=0.87)
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Partly supported
The cited sources that could be checked couldn’t confirm or dispute this claim.
In the paper
“Natural language models often struggle to maintain coherent and diverse generation over long sequences”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
What the sources say
- Could not verifycites [39] Kaplan J., Scaling Laws for Neural Language Models. arXiv: 2001. 08361 [cs.LG] (2020).Experimental — accuracy not yet measured
The Introduction (p. 2) says recent models can compose coherent multiparagraph text, while Appendix D.5 (p. 25) examines loss and longer-range correlations across a context. Neither passage addresses whether language models often struggle to maintain coherence and diversity when generating long sequences; searches for those terms found no supporting passage. (LLM, conf=0.76)
- Could not verifycites [83] Wei J., Finetuned language models are zero-shot learners. arXiv: 2109. 01652 [cs.CL] (2022). sourceExperimental — accuracy not yet measured
The Introduction (p. 2) discusses difficulty with zero-shot tasks, not maintaining coherence or diversity in long text. The qualitative examples (p. 34) include a story of at least 150 tokens but do not make the claimed general statement. Searches for coherence, diversity, and long-form generation found no passage addressing it. (LLM, conf=0.76)
- Could not verifycites [84] Ouyang L., Training language models to follow instructions with human feedback. arXiv: 2203. 02155 [cs.CL] (2022). sourceExperimental — accuracy not yet measured
The Introduction describes failures such as making up facts, producing biased or toxic text, and not following instructions, but does not discuss coherence or diversity over long sequences. Searches for long-form generation, repetition, and output diversity found no passage establishing the claimed difficulty; the References merely list a work on coherent long-form text generation. (LLM, conf=0.72)
- Could not verifycites [85] Rafailov R., “ Direct Preference Optimization: Your Language Model is Secretly a Reward Model” in Advances in Neural Information Processing Systems, vol. 36, Oh A. et al., Eds. (Curran Associates, Inc., 2023), pp. 53728– 53741. sourceExperimental — accuracy not yet measured
The Additional information section says that an unlikelihood-training baseline produced meaningless responses on summarization and dialogue tasks, but it does not discuss whether language models generally struggle with coherence and diversity over long sequences. Searches for coherent and diverse long-sequence generation found no passage addressing that claim; the copy provides no page number for the cited section. (LLM, conf=0.72)
- Could not verifycites [87] Kojima T, Gu SS, Reid M, Matsuo Y, Iwasawa Y, “ Large Language Models are Zero-Shot Reasoners” in Advances in Neural Information Processing Systems, vol. 35, Koyejo S., Eds. (Curran Associates, Inc., 2022), pp. 22199– 22213. sourceExperimental — accuracy not yet measured
The discussion of Few-shot-CoT (p. 2) says language models have struggled with multi-step reasoning, not with maintaining coherence or diversity in long text generation. Searches for long-sequence generation, coherence, diversity, and repetition found no passage in our copy addressing the attributed claim. (LLM, conf=0.76)
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Could not verify
The cited sources support only part of this claim — read each source’s verdict below.
In the paper
“Multistage sequence length pretraining has been shown to reduce the overall number of compute hours required to train long context models (116).”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Training procedure
What the sources say
- Partly supportedcites [116] Xiong W., Effective long-context scaling of foundation models. arXiv: 2309. 16039 [cs.CL] (2023). sourceExperimental — accuracy not yet measured
Section 4.4 (Training Curriculum; page not available in our copy) compares two-stage sequence-length training with training at the long sequence length throughout and reports about 40% fewer FLOPs with almost no performance loss. That supports reduced computational work, but the paper does not report compute hours, so the claim's specific time-based measure is not established. (LLM, conf=0.82)
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Partly supported
Every source the paper cites for this claim was checked and supports it.
In the paper
“We experimented with grouped-query attention (GQA) (117) and found minimal differences in final loss, suggesting the technique may be suited to DNA sequence modeling, to further reduce memory footprint during inference.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Transformer++
What the sources say
- Supportedcites [117] Ainslie J., GQA: Training generalized multi-query transformer models from multi-head checkpoints. arXiv: 2305. 13245 [cs.CL] (2023). source
The citation identifies grouped-query attention, which Section 2.2 defines as sharing key and value heads across groups of query heads and describes as reducing inference memory demands. The citing authors present the minimal change in final loss and the possible application to DNA modeling as findings and inferences from their own experiment, not as findings attributed to this source. (LLM, conf=0.93)
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
- Supported
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Evo is a genomic foundation model with 7 billion parameters trained with a context length of up to 131,072 tokens, using single-nucleotide, byte-level tokenization.”
§ Sequence modeling and design from molecular to genome scale with Evo > Modeling long sequences at nucleotide resolution with the StripedHyena architecture
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We estimate 250 billion to be the compute-optimal number of tokens for Evo 7B given the floating point operation (FLOP) budget, meaning the model was trained at a 17% offset from the compute-optimal model size during the initial 8192 sequence length pretraining phase of 300 billion tokens.”
§ Sequence modeling and design from molecular to genome scale with Evo > StripedHyena demonstrates favorable scaling laws on DNA sequence data
hedged- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Both Transformer++ and Mamba experienced numerical instability during training and suffered from a higher performance degradation of the scaling rate outside the compute-optimal frontier, in contrast to StripedHyena”
§ Sequence modeling and design from molecular to genome scale with Evo > StripedHyena demonstrates favorable scaling laws on DNA sequence data
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Evo uses a context length of 131,072 tokens and is based on the StripedHyena architecture (24), which hybridizes attention and data-controlled convolutional operators to efficiently process and recall patterns in long sequences.”
§ Sequence modeling and design from molecular to genome scale with Evo > Abstract > CONCLUSION:
What the sources say
- Not checked yetcites [24] Poli M., StripedHyena: Moving Beyond Transformers with Hybrid Signal Processing Models, GitHub (2023); https://github.com/togethercomputer/stripedhyena.
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“One of the generations exhibited robust activity, which we named EvoCas9–1.”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We observed that more conserved sequences tended to be essential, with an association strength similar to that of Evo with gene-only context but weaker than that of Evo with genomic sequence context”
§ Sequence modeling and design from molecular to genome scale with Evo > Learning gene essentiality with long genomic context
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Although the predicted backbone structure of EvoCas9–1 resembles that of SpCas9, the predicted structure of EvoCas9–1 exhibits a more positive surface charge distribution”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“when concatenating the promoter and RBS sequence together, Evo’s zero-shot likelihoods improved substantially (Spearman r = 0.61); this correlation is also higher than the zero-shot correlation of just the promoter sequence alone (Spearman r = 0.47)”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting activity of regulatory DNA
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“TnpA and TnpB proteins that were detected within these generated sequences varied widely in their distance from the nearest examples in the training set (Fig. 4C), with consistently high ESMFold pLDDT values for predicted structures that were >40 to 50% identity to the training set”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of transposon systems
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one. The quote only partly states this claim.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This activity was also dependent on the presence of a putative catalytic tyrosine and on having a ssDNA substrate instead of double-stranded DNA (dsDNA), consistent with the known mechanism for IS200/IS605 TnpA”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of transposon systems
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These correlations also exceed those of the sequence guanine-cytosine (GC) content (mean Spearman r = 0.35) and the zero-shot likelihoods of GenSLM (mean Spearman r = 0.09)”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting activity of regulatory DNA
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Evo-designed sgRNA is 91.1% identical to the canonical SpCas9 sgRNA and exhibits secondary structure differences in the two terminal stem loops, notably extending the length of stem loops 2 and 3”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We observed that 11 out of 24 Evo-generated IS200-like elements and 3 out of 24 Evo-generated IS605-like elements demonstrated evidence for both excision and insertion in vitro”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of transposon systems
boosted- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The EvoCas9–1 amino acid sequence shares 79.9% identity with the closest Cas9 in the database of Cas proteins used for model fine-tuning and 73.1% identity with SpCas9.”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We also observed that providing the model with additional genomic context beyond the gene sequence results in a substantial improvement in performance, especially from gene-only context to 8k context. From 8k to 66k context, the average predictive performance is comparable, although performance on the lower range of examples does improve with longer context”
§ Sequence modeling and design from molecular to genome scale with Evo > Learning gene essentiality with long genomic context
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“As a whole, the 14 active elements use a diverse set of hairpins (Fig. 4, E and H, and figs. S25 and S26) and encode functional TnpA proteins with sequence identity as low as 67% to the fine-tuning database.”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of transposon systems
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The CNN architecture substantially outperformed ridge regression across both embeddings, and the Evo embeddings substantially outperformed one-hot embeddings across both architectures”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting activity of regulatory DNA
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The functional IS605-like elements, which contain putative TnpB coding sequences, also contain sequences with significant matches (cmsearch E value < 0.001) to a covariance model constructed from known ωRNAs”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of transposon systems
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A claim is checked only against the cited paper’s full text, never its abstract, and none is available for its sources.
In the paper
“Compared with HyenaDNA (33), a previous generation of DNA models leveraging a Hyena architecture (34), Evo is based on an improved hybrid design and scaled to 1000× larger model size and 100× more data.”
§ Sequence modeling and design from molecular to genome scale with Evo > Modeling long sequences at nucleotide resolution with the StripedHyena architecture
What the sources say
- Not checked — no full text of the sourcecites [33] Nguyen E., “ HyenaDNA: Long-Range Genomic Sequence Modeling at Single Nucleotide Resolution” in Advances in Neural Information Processing Systems, vol. 36, Oh A. et al., Eds. (Curran Associates, Inc., 2023), pp. 43177– 43201. source
- Not checked — no full text of the sourcecites [34] Poli M., Hyena Hierarchy: Towards Larger Convolutional Language Models. arXiv: 2302. 10866 [cs.LG] (2023). source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Evo is a genomic foundation model trained on hundreds of billions of DNA tokens across the evolutionary diversity of prokaryotic life, capable of prediction and generation tasks at the scale of individual molecules, molecular complexes, systems, and even whole genomes.”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“When visualized, both natural and generated sequences display similar patterns of coding organization (Fig. 6C), with sequences in close proximity typically found with the same strand orientation”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Based on a state-of-the art hybrid model architecture, Evo enables single-nucleotide-resolution language modeling at a context length of 131,072.”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We used Evo to sample 16 sequences each containing ~1 Mb, representing more than seven times the model’s context length of 131 kb.”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Evo accurately performed zero-shot prediction across diverse fitness or expression prediction tasks on proteins, ncRNAs, or regulatory DNA that matches or outperforms specialized models while also understanding how mutations to individual genes can affect broader organismal fitness.”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“As a multimodal generative model, we use Evo to generate CRISPR-Cas proteins and their noncoding guide RNAs, multicomponent transposable systems, and megabase-long sequences that recapitulate the architecture of real genomes.”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We found strong correlations between species-specific generations and their corresponding natural reference sequences, with TUDs sufficiently accurate to reconstruct natural phylogenetic relationships among the generated sequences (Fig. 6, F and G).”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
boostedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: TUDs computed on Evo-generated sequences can reconstruct phylogenetic relationships among those sequences.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We conducted the first scaling laws analysis of DNA pretraining across several architectures, where we observed StripedHyena outperforming several baseline architectures, including Transformers.”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Notably, Evo generated sequences have nearly the same coding densities as natural genomes, and substantially higher than that of random sequences (Fig. 6B).”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: CheckM metrics computed on Evo-generated sequences are comparable to those computed on natural bacterial genomes.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Evo can demonstrate similar properties.”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The quote only partly states this claim.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We pretrained Evo on a dataset of 300 billion prokaryotic tokens, which represents a miniscule portion of petabytes of publicly available genomic data.”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“At the genome-scale, Evo generates megabase-long sequences that demonstrate a high-level understanding of genome organization, but it struggles to include key marker genes, such as full sets of rRNAs.”
§ Sequence modeling and design from molecular to genome scale with Evo > Discussion
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“we found that, in some instances, adding an additional random nucleotide character would improve the quality of generations.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > CRISPR-Cas fine-tuning and generation
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We experimentally validated the functional activity of Evo-generated CRISPR-Cas molecular complexes as well as IS200 and IS605 transposable systems”
§ Sequence modeling and design from molecular to genome scale with Evo > Abstract > RESULTS:
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“EvoCas9–1 was generated from just 11 code-signs, representing a robust success rate given the complexity of Cas9’s multistep mechanism”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
plainSingle case- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Model hybridization, first proposed to address shortcomings of state-space models (30–32), has recently been shown to improve scaling performance on language modeling of both standalone Hyena and Transformer architectures (24).”
§ Sequence modeling and design from molecular to genome scale with Evo > Modeling long sequences at nucleotide resolution with the StripedHyena architecture
What the sources say
- Not checked yetcites [24] Poli M., StripedHyena: Moving Beyond Transformers with Hybrid Signal Processing Models, GitHub (2023); https://github.com/togethercomputer/stripedhyena.
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Evo is a foundation model that is designed to capture two fundamental aspects of biology: the multimodality of the central dogma and the multiscale nature of evolution.”
§ Sequence modeling and design from molecular to genome scale with Evo > Abstract > CONCLUSION:
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Because Evo learns covariation patterns involving any genetic elements within its context window, the model should understand interactions between encoded protein and ncRNA molecules.”
§ Sequence modeling and design from molecular to genome scale with Evo > Generative design of CRISPR-Cas molecular complexes
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper may give no reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Evo demonstrates a robust and general understanding of the constitutive protein coding sequences, ncRNA sequences, and regulatory elements.”
§ Sequence modeling and design from molecular to genome scale with Evo > Evo learns across DNA, RNA, and protein modalities > Predicting activity of regulatory DNA
boostedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“For comparison, the smallest “minimal” bacterial genomes are ~580 kb in length (76).”
§ Sequence modeling and design from molecular to genome scale with Evo > Generating DNA sequences at genome scale
What the sources say
- Not checked — no full text of the sourcecites [76] Blanchard A, Bébéar C, The evolution of Mycoplasma genitalium. Ann. N. Y. Acad. Sci 1230, E61– E64 (2011). doi: 10.1111/j.1749-6632.2011.06418.x 22417108 source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“As a negative control, we generated 1000 sequences of length 131,072 in which the four DNA base pairs were sampled uniformly at random.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Genome-scale generation and evaluation
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“To account for different start codons and to generate a more conservative percentage identity estimate, these alignments were trimmed to the middle 80% of each sequence, end gaps were trimmed, and the amino acid percent identity was recalculated, which we called a “trimmed percent identity.””
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > IS200/IS605 fine-tuning and generation
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“CheckM uses Prodigal (140) to identify coding sequences and computes the coding density as one metric of genome quality.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Genome-scale generation and evaluation
What the sources say
- Not matchedcites CheckM (77)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Not matched
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“To select candidates for experimental validation, Cas9 generations with an ORF sequence identity higher than 90% to a training sequence were first filtered out.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > CRISPR-Cas sampling evaluation
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“As an additional control, we also used the gene’s linear position in the reference genome as the value with which to predict essentiality.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Gene essentiality prediction
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“We evaluated these generations with CheckM (77), a tool that computes basic genome quality metrics based on whether a given long DNA sequence has similar properties as known bacterial genomes.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Genome-scale generation and evaluation
What the sources say
- Not matchedcites CheckM (77)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
- Not matched
In the paper
“CheckM will also search for the presence of genes that are highly conserved across much of prokaryotic diversity.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Genome-scale generation and evaluation
What the sources say
- Not matchedcites CheckM (77)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Not matched
None of the sources cited for this claim could be matched to a paper in Crossref, so there is nothing to check it against.
In the paper
“CheckM uses Prodigal (140) to identify coding sequences and computes the coding density as one metric of genome quality.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Genome-scale generation and evaluation
What the sources say
- Not matchedcites CheckM (77)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Not matchedcites Prodigal (140)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Not matched
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“For final synthesis and experimental validation, a different 60 bp filler sequence was used for the ISHp608-like candidates compared to the ISStin10-like, ISSpn6-like, or ISDge10-like sequences to eliminate a primer-binding site containing a TTAC, which is the canonical ISHp608 target site.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > IS200/IS605 filtering of generations and construct design
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“We assessed statistical significance of the AUROC with a permutation-based method in which a null distribution is constructed by permuting the binary labels and recomputing the subsequent AUROC.”
- Also stated in: § Gene essentiality prediction · plain
“We performed 100,000 permutations to construct this null distribution.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Gene essentiality prediction
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Also stated in: § Gene essentiality prediction · plain
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“To ensure that the Spearman correlation is comparable across these settings, we computed the correlation over all 12,243 examples (which involves duplicating sequences in the promoter-alone or RBS-alone settings).”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Gene expression prediction from regulatory DNA
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“We also used the conservation of a gene as another control.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Gene essentiality prediction
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- The authors state this as an assumption: A greater number of significant mmseqs hits corresponds to higher conservation.
- The authors state this as an assumption: Higher conservation corresponds to greater essentiality.
None of the sources cited for this claim could be matched to a paper in Crossref, so there is nothing to check it against.
In the paper
“As a positive control, we randomly selected 100 bacterial genomes from GTDB and used CheckM to compute the coding densities for 131,072 bp crops from these genomes.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Genome-scale generation and evaluation
What the sources say
- Not matchedcites GTDB
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Not matchedcites CheckM (77)
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Not matched
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“We also modified the data loader such that each sample provided to the model during training would begin with the first token of the IS200/IS605 sequence and, if a sequence was shorter than the context length, we padded the sequence to the remaining context (where padding did not contribute to the loss computation), similar to the strategy described for CRISPR-Cas9 systems above.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > IS200/IS605 fine-tuning and generation
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“If a model were simply using positional information to make essentiality predictions, the performance would be similar to this control.”
§ Sequence modeling and design from molecular to genome scale with Evo > Materials and methods > Gene essentiality prediction
hedged- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
The paper relies on this without support.
Functionally related genes in prokaryotes are generally organized into operons and located next to each other on the genome.
“In prokaryotes, functionally related genes are generally organized into operons and located next to each other on the genome sequence.”
The paper relies on this without support.
Transcription from a CRISPR array produces crRNA that is bound by a Cas protein to form a functional defense complex required for sequence-specific DNA targeting.
“Transcription from the CRISPR array generates noncoding CRISPR RNA (crRNA) molecules that are bound by the Cas protein to generate a functional defense complex that is required for sequence-specific DNA targeting”
The paper relies on this without support.
For Cas9, a tracrRNA forms a duplex with the crRNA to create a full guide RNA.
“For Cas9 in particular, a second trans-activating CRISPR RNA (tracrRNA) forms a duplex with the crRNA to create a full guide RNA (gRNA).”
The paper relies on this without support.
Successful transposition requires TnpA proteins to dimerize, interact with DNA hairpins at the left and right ends, base-pair with the target site, and carry out strand cleavage and exchange.
“successful transposition requires TnpA proteins that functionally dimerize, TnpA dimer interactions with DNA hairpins in the LE and RE, base pairing between the LE and RE hairpins and the target site, and strand cleavage and exchange.”
The paper relies on this without support.
In vitro cleavage, excision, and insertion activity is a valid proxy for the biological function of Evo-generated CRISPR-Cas and transposon systems.
“We experimentally validated the functional activity of Evo-generated CRISPR-Cas molecular complexes as well as IS200 and IS605 transposable systems”
The paper relies on this without support.
In bacteria, adjacent coding sequences on the same strand often form operons or gene clusters with related functions.
“in bacteria, these closely linked groups of coding sequences typically correspond to functionally tied gene clusters or operons.”
The paper relies on this without support.
CheckM metrics computed on Evo-generated sequences are comparable to those computed on natural bacterial genomes.
“we used CheckM (77), a tool originally designed to assess the quality of bacterial DNA sequenced from nature.”
The paper relies on this without support.
TUDs computed on Evo-generated sequences can reconstruct phylogenetic relationships among those sequences.
“we calculated tetranucleotide usage deviations (TUDs), a strong indicator metric of phylogenetic relatedness (78).”
The authors state this as an assumption.
A greater number of significant mmseqs hits corresponds to higher conservation.
“We assumed that a greater number of hits corresponds to higher conservation, which in turn corresponds to greater essentiality.”
The authors state this as an assumption.
Higher conservation corresponds to greater essentiality.
“We assumed that a greater number of hits corresponds to higher conservation, which in turn corresponds to greater essentiality.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The cited source says what the claim attributes to it.
In the citing paper
“AI models have already been used to design new CRISPR enzymes,53 functional proteins,128 and even entire prokaryotic genomes.51”
From How to build the virtual cell with artificial intelligence: Priorities and opportunities (2024) · cites this paper as [51]
Citations
Cites
Basic local alignment search tool
Altschul, S. F., Gish, W., Miller, W., Myers, E. W., & Lipman, D. J. (1990) · Journal of Molecular Biology · doi:10.1006/jmbi.1990.9999 · Cited by 98,669
Highly accurate protein structure prediction with AlphaFold
Jumper, J., Evans, R., Pritzel, A., Green, T., Figurnov, M., Ronneberger, O., Tunyasuvunakool, K., Bates, R., Žídek, A., Potapenko, A., et al. (2021) · Nature · doi:10.1038/s41586-021-03819-2 · Cited by 43,764
A Programmable Dual-RNA–Guided DNA Endonuclease in Adaptive Bacterial Immunity
Jinek, M., Chylinski, K., Fonfara, I., Hauer, M., Doudna, J. A., & Charpentier, E. (2012) · Science · doi:10.1126/science.1225829 · Cited by 17,116
MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform
Katoh, K. (2002) · Nucleic Acids Research · doi:10.1093/nar/gkf436 · Cited by 17,027
Advances in Neural Information Processing Systems 19
(2007) · The MIT Press eBooks · doi:10.52202/085713 · Cited by 16,043
Accurate structure prediction of biomolecular interactions with AlphaFold 3
Abramson, J., Adler, J., Dunger, J., Evans, R., Green, T., Pritzel, A., Ronneberger, O., Willmore, L., Ballard, A. J., Bambrick, J., et al. (2024) · Nature · doi:10.1038/s41586-024-07487-w · Cited by 15,861
Molecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid
WATSON, J. D., & CRICK, F. H. C. (1953) · Nature · doi:10.1038/171737a0 · Cited by 13,584
Prodigal: prokaryotic gene recognition and translation initiation site identification
Hyatt, D., Chen, G.-L., LoCascio, P. F., Land, M., Larimer, F. W., & Hauser, L. (2010) · BMC Bioinformatics · doi:10.1186/1471-2105-11-119 · Cited by 12,692
Parks, D. H., Imelfort, M., Skennerton, C. T., Hugenholtz, P., & Tyson, G. W. (2015) · Genome Research · doi:10.1101/gr.186072.114 · Cited by 12,496
Proceedings of the 24th international conference on Machine learning
(2007) · doi:10.1145/1273496 · Cited by 11,869
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 · cited 2×
“Aidingourmodeldesign,weperformedascaling laws analysis for DNA sequence modeling to determine the relationship between training, architectural details, and performance metrics through a systematic experimental protocol ( 38 , 39 ).”
ColabFold - Making protein folding accessible to all
Mirdita, M., Schütze, K., Moriwaki, Y., Heo, L., Ovchinnikov, S., & Steinegger, M. (2021) · doi:10.1101/2021.08.15.456425 · Cited by 9,391
O'Leary, N. A., Wright, M. W., Brister, J. R., Ciufo, S., Haddad, D., McVeigh, R., Rajput, B., Robbertse, B., Smith-White, B., Ako-Adjei, D., et al. (2016) · Nucleic Acids Research · doi:10.1093/nar/gkv1189 · Cited by 6,357
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
Self-citationHMMER web server: interactive sequence similarity searching
Finn, R. D., Clements, J., & Eddy, S. R. (2011) · Nucleic Acids Research · doi:10.1093/nar/gkr367 · Cited by 5,952
Development and Applications of CRISPR-Cas9 for Genome Engineering
Hsu, P. D., Lander, E. S., & Zhang, F. (2014) · Cell · doi:10.1016/j.cell.2014.05.010 · Cited by 5,762
Self-citationLorenz, R., Bernhart, S. H., Höner zu Siederdissen, C., Tafer, H., Flamm, C., Stadler, P. F., & Hofacker, I. L. (2011) · Algorithms for Molecular Biology · doi:10.1186/1748-7188-6-26 · Cited by 5,098
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
DNA targeting specificity of RNA-guided Cas9 nucleases
Hsu, P. D., Scott, D. A., Weinstein, J. A., Ran, F. A., Konermann, S., Agarwala, V., Li, Y., Fine, E. J., Wu, X., Shalem, O., et al. (2013) · Nature Biotechnology · doi:10.1038/nbt.2647 · Cited by 4,741
Self-citationTraining language models to follow instructions with human feedback
Ouyang, L., Wu, J., Jiang, X., Almeida, D., Wainwright, C. L., Mishkin, P., Zhang, C., Agarwal, S., Slama, K., Ray, A., et al. (2022) · arXiv · doi:10.48550/arxiv.2203.02155 · Cited by 4,208
Fourier Neural Operator for Parametric Partial Differential EquationsPreprint
Li, Z., Kovachki, N., Azizzadenesheli, K., Liu, B., Bhattacharya, K., Stuart, A. M., & Anandkumar, A. (2020) · arXiv (Cornell University) · doi:10.48550/arxiv.2010.08895 · Cited by 3,717
Infernal 1.1: 100-fold faster RNA homology searches
Nawrocki, E. P., & Eddy, S. R. (2013) · Bioinformatics · doi:10.1093/bioinformatics/btt509 · Cited by 3,688
Meyer, F., Paarmann, D., D'Souza, M., Olson, R., Glass, E., Kubal, M., Paczian, T., Rodriguez, A., Stevens, R. L., Wilke, A., et al. (2008) · BMC Bioinformatics · doi:10.1186/1471-2105-9-386 · Cited by 3,565
Rives, A., Meier, J., Sercu, T., Goyal, S., Lin, Z., Liu, J., Guo, D., Ott, M., Zitnick, C. L., Ma, J., et al. (2021) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2016239118 · Cited by 3,286
Rives, A., Meier, J., Sercu, T., Goyal, S., Lin, Z., Liu, J., Guo, D., Ott, M., Zitnick, C. L., Ma, J., et al. (2019) · doi:10.1101/622803 · Cited by 3,280
Training Language Models to Follow Instructions with Human Feedback
Ouyang, L., Wu, J., Jiang, X., Almeida, D., Wainwright, C., Mishkin, P., Zhang, C., Agarwal, S., Slama, K., Ray, A., et al. (2022) · Advances in Neural Information Processing Systems 35 · doi:10.52202/068431-2011 · Cited by 3,117
Structure and function of the global ocean microbiome
Sunagawa, S., Coelho, L. P., Chaffron, S., Kultima, J. R., Labadie, K., Salazar, G., Djahanschiri, B., Zeller, G., Mende, D. R., Alberti, A., et al. (2015) · Science · doi:10.1126/science.1261359 · Cited by 2,978
ISfinder: the reference centre for bacterial insertion sequences
Siguier, P. (2006) · Nucleic Acids Research · doi:10.1093/nar/gkj014 · Cited by 2,767
Gruber, A. R., Lorenz, R., Bernhart, S. H., Neuböck, R., & Hofacker, I. L. (2008) · Nucleic Acids Research · doi:10.1093/nar/gkn188 · Cited by 2,717
Parks, D. H., Chuvochina, M., Rinke, C., Mussig, A. J., Chaumeil, P.-A., & Hugenholtz, P. (2022) · Nucleic Acids Research · doi:10.1093/nar/gkab776 · Cited by 2,402
Watson, J. L., Juergens, D., Bennett, N. R., Trippe, B. L., Yim, J., Eisenach, H. E., Ahern, W., Borst, A. J., Ragotte, R. J., Milles, L. F., et al. (2022) · doi:10.1101/2022.12.09.519842 · Cited by 2,369
De novo design of protein structure and function with RFdiffusion
Watson, J. L., Juergens, D., Bennett, N. R., Trippe, B. L., Yim, J., Eisenach, H. E., Ahern, W., Borst, A. J., Ragotte, R. J., Milles, L. F., et al. (2023) · Nature · doi:10.1038/s41586-023-06415-8 · Cited by 2,325
Large language models are zero-shot reasoners
Kojima, T., Gu, S. (., Reid, M., Matsuo, Y., & Iwasawa, Y. (2022) · Advances in Neural Information Processing Systems 35 · doi:10.52202/068431-1613 · Cited by 2,013
RoFormer: Enhanced transformer with rotary position embedding
Su, J., Ahmed, M., Lu, Y., Pan, S., Bo, W., & Liu, Y. (2024) · Neurocomputing · doi:10.1016/j.neucom.2023.127063 · Cited by 1,957
Automated design of synthetic ribosome binding sites to control protein expression
Salis, H. M., Mirsky, E. A., & Voigt, C. A. (2009) · Nature Biotechnology · doi:10.1038/nbt.1568 · Cited by 1,844
tRNAscan-SE: Searching for tRNA Genes in Genomic Sequences
Chan, P. P., & Lowe, T. M. (2019) · Methods in molecular biology · doi:10.1007/978-1-4939-9173-0_1 · Cited by 1,821
Language Modeling with Gated Convolutional NetworksPreprint
Dauphin, Y., Fan, A., Auli, M., & Grangier, D. (2016) · arXiv (Cornell University) · doi:10.48550/arxiv.1612.08083 · Cited by 1,765
Nirenberg, M. W., & Matthaei, J. (1961) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.47.10.1588 · Cited by 1,638
Mamba: Linear-Time Sequence Modeling with Selective State Spaces
Gu, A., & Dao, T. (2023) · arXiv · doi:10.48550/arxiv.2312.00752 · Cited by 1,579
A unified catalog of 204,938 reference genomes from the human gut microbiome
Almeida, A., Nayfach, S., Boland, M., Strozzi, F., Beracochea, M., Shi, Z. J., Pollard, K. S., Sakharova, E., Parks, D. H., Hugenholtz, P., et al. (2021) · Nature Biotechnology · doi:10.1038/s41587-020-0603-3 · Cited by 1,367
Effective gene expression prediction from sequence by integrating long-range interactions
Avsec, Ž., Agarwal, V., Visentin, D., Ledsam, J. R., Grabska-Barwinska, A., Taylor, K. R., Assael, Y., Jumper, J., Kohli, P., & Kelley, D. R. (2021) · Nature Methods · doi:10.1038/s41592-021-01252-x · Cited by 1,326
Ji, Y., Zhou, Z., Liu, H., & Davuluri, R. V. (2021) · Bioinformatics · doi:10.1093/bioinformatics/btab083 · Cited by 1,303
Genetics and the Origin of Species
Dobzhansky, T. (1952) · Copeia · doi:10.2307/1439305 · Cited by 1,265
Transfer learning enables predictions in network biology
Theodoris, C. V., Xiao, L., Chopra, A., Chaffin, M. D., Al Sayed, Z. R., Hill, M. C., Mantineo, H., Brydon, E. M., Zeng, Z., Liu, X. S., et al. (2023) · Nature · doi:10.1038/s41586-023-06139-9 · Cited by 1,193
Finetuned Language Models Are Zero-Shot Learners
Wei, J., Bosma, M., Zhao, V. Y., Guu, K., Yu, A. W., Lester, B., Du, N., Dai, A. M., & Le, Q. V. (2021) · arXiv · doi:10.48550/arxiv.2109.01652 · Cited by 1,101
Large language models generate functional protein sequences across diverse families
Madani, A., Krause, B., Greene, E. R., Subramanian, S., Mohr, B. P., Holton, J. M., Olmos, J. L., Xiong, C., Sun, Z. Z., Socher, R., et al. (2023) · Nature Biotechnology · doi:10.1038/s41587-022-01618-2 · Cited by 1,078
Bland, C., Ramsey, T. L., Sabree, F., Lowe, M., Brown, K., Kyrpides, N. C., & Hugenholtz, P. (2007) · BMC Bioinformatics · doi:10.1186/1471-2105-8-209 · Cited by 1,038
Training Compute-Optimal Large Language Models
Hoffmann, J., Borgeaud, S., Mensch, A., Buchatskaya, E., Cai, T., Rutherford, E., Casas, D. D. L., Hendricks, L. A., Welbl, J., Clark, A., et al. (2022) · Advances in Neural Information Processing Systems 35 · doi:10.52202/068431-2176 · Cited by 932
Training Compute-Optimal Large Language Models
Hoffmann, J., Borgeaud, S., Mensch, A., Buchatskaya, E., Cai, T., Rutherford, E., Casas, D. D. L., Hendricks, L. A., Welbl, J., Clark, A., et al. (2022) · arXiv · doi:10.48550/arxiv.2203.15556 · Cited by 929
Accurate classification of BRCA1 variants with saturation genome editing
Findlay, G. M., Daza, R. M., Martin, B., Zhang, M. D., Leith, A. P., Gasperini, M., Janizek, J. D., Huang, X., Starita, L. M., & Shendure, J. (2018) · Nature · doi:10.1038/s41586-018-0461-z · Cited by 843
Efficiently Modeling Long Sequences with Structured State Spaces
Gu, A., Goel, K., & Ré, C. (2021) · arXiv · doi:10.48550/arxiv.2111.00396 · Cited by 805
Self-citationBLAST: improvements for better sequence analysis
Ye, J., McGinnis, S., & Madden, T. (2006) · Nucleic Acids Research · doi:10.1093/nar/gkl164 · Cited by 774
Nucleotide Transformer: building and evaluating robust foundation models for human genomics
Dalla-Torre, H., Gonzalez, L., Mendoza-Revilla, J., Lopez Carranza, N., Grzywaczewski, A. H., Oteri, F., Dallago, C., Trop, E., de Almeida, B. P., Sirelkhatim, H., et al. (2025) · Nature Methods · doi:10.1038/s41592-024-02523-z · Cited by 772
Direct Preference Optimization: Your Language Model is Secretly a Reward Model
Rafailov, R., Sharma, A., Mitchell, E., Manning, C. D., Ermon, S., & Finn, C. (2023) · Advances in Neural Information Processing Systems 36 · doi:10.52202/075280-2338 · Cited by 686
Self-citationThe Nucleotide Transformer: Building and Evaluating Robust Foundation Models for Human Genomics
Dalla-Torre, H., Gonzalez, L., Mendoza Revilla, J., Lopez Carranza, N., Henryk Grywaczewski, A., Oteri, F., Dallago, C., Trop, E., De Almeida, B. P., Sirelkhatim, H., et al. (2023) · doi:10.1101/2023.01.11.523679 · Cited by 680
Language models enable zero-shot prediction of the effects of mutations on protein function
Meier, J., Rao, R., Verkuil, R., Liu, J., Sercu, T., & Rives, A. (2021) · doi:10.1101/2021.07.09.450648 · Cited by 635
Sex Limited Inheritance in Drosophila
Morgan, T. H. (1910) · Science · doi:10.1126/science.32.812.120 · Cited by 619
Massive expansion of human gut bacteriophage diversity
Camarillo-Guerrero, L. F., Almeida, A., Rangel-Pineros, G., Finn, R. D., & Lawley, T. D. (2021) · Cell · doi:10.1016/j.cell.2021.01.029 · Cited by 594
DEG: a database of essential genes
Zhang, R. (2003) · Nucleic Acids Research · doi:10.1093/nar/gkh024 · Cited by 572
A human gut bacterial genome and culture collection for improved metagenomic analyses
Forster, S. C., Kumar, N., Anonye, B. O., Almeida, A., Viciani, E., Stares, M., Dunn, M., Mkandawire, T. T., Zhu, A., Shao, Y., et al. (2019) · Nature Biotechnology · doi:10.1038/s41587-018-0009-7 · Cited by 566
MGnify: the microbiome analysis resource in 2020
Mitchell, A. L., Almeida, A., Beracochea, M., Boland, M., Burgin, J., Cochrane, G., Crusoe, M. R., Kale, V., Potter, S. C., Richardson, L. J., et al. (2019) · Nucleic Acids Research · doi:10.1093/nar/gkz1035 · Cited by 562
Mutational processes shape the landscape of TP53 mutations in human cancer
Giacomelli, A. O., Yang, X., Lintner, R. E., McFarland, J. M., Duby, M., Kim, J., Howard, T. P., Takeda, D. Y., Ly, S. H., Kim, E., et al. (2018) · Nature Genetics · doi:10.1038/s41588-018-0204-y · Cited by 556
US-align: universal structure alignments of proteins, nucleic acids, and macromolecular complexes
Zhang, C., Shine, M., Pyle, A. M., & Zhang, Y. (2022) · Nature Methods · doi:10.1038/s41592-022-01585-1 · Cited by 511
Origins and evolution of CRISPR-Cas systems
Koonin, E. V., & Makarova, K. S. (2019) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2018.0087 · Cited by 494
ProGen2: Exploring the boundaries of protein language models
Nijkamp, E., Ruffolo, J. A., Weinstein, E. N., Naik, N., & Madani, A. (2023) · Cell Systems · doi:10.1016/j.cels.2023.10.002 · Cited by 480
Efficient evolution of human antibodies from general protein language models
Hie, B., Shanker, V. R., Xu, D., Bruun, T. U. J., Weidenbacher, P. A. B., Tang, S., Wu, W. C. H., Pak, J. E., & Kim, P. S. (2023) · Nature Biotechnology · doi:10.1038/s41587-023-01763-2 · Cited by 472
Self-citationGQA: Training Generalized Multi-Query Transformer Models from Multi-Head Checkpoints
Ainslie, J., Lee-Thorp, J., de Jong, M., Zemlyanskiy, Y., Lebron, F., & Sanghai, S. (2023) · Proceedings of the 2023 Conference on Empirical Methods in Natural Language Processing · doi:10.18653/v1/2023.emnlp-main.298 · Cited by 454
GQA: Training Generalized Multi-Query Transformer Models from Multi-Head Checkpoints
Ainslie, J., Lee-Thorp, J., de Jong, M., Zemlyanskiy, Y., Lebrón, F., & Sanghai, S. (2023) · arXiv · doi:10.48550/arxiv.2305.13245 · Cited by 447
Camargo, A. P., Nayfach, S., Chen, I.-M. A., Palaniappan, K., Ratner, A., Chu, K., Ritter, S. J., Reddy, T. B. K., Mukherjee, S., Schulz, F., et al. (2023) · Nucleic Acids Research · doi:10.1093/nar/gkac1037 · Cited by 446
Illuminating protein space with a programmable generative model
Ingraham, J. B., Baranov, M., Costello, Z., Barber, K. W., Wang, W., Ismail, A., Frappier, V., Lord, D. M., Ng-Thow-Hing, C., Van Vlack, E. R., et al. (2023) · Nature · doi:10.1038/s41586-023-06728-8 · Cited by 444
The IMG/M data management and analysis system v.6.0: new tools and advanced capabilities
Chen, I.-M. A., Chu, K., Palaniappan, K., Ratner, A., Huang, J., Huntemann, M., Hajek, P. P., Ritter, S., Varghese, N. J., Seshadri, R., et al. (2020) · Nucleic Acids Research · doi:10.1093/nar/gkaa939 · Cited by 433
Essential genome of Pseudomonas aeruginosa in cystic fibrosis sputum
Turner, K. H., Wessel, A. K., Palmer, G. C., Murray, J. L., & Whiteley, M. (2015) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1419677112 · Cited by 432
The widespread IS200/IS605 transposon family encodes diverse programmable RNA-guided endonucleases
Altae-Tran, H., Kannan, S., Demircioglu, F. E., Oshiro, R., Nety, S. P., McKay, L. J., Dlakić, M., Inskeep, W. P., Makarova, K. S., Macrae, R. K., et al. (2021) · Science · doi:10.1126/science.abj6856 · Cited by 407
Composability of regulatory sequences controlling transcription and translation in Escherichia coli
Kosuri, S., Goodman, D. B., Cambray, G., Mutalik, V. K., Gao, Y., Arkin, A. P., Endy, D., & Church, G. M. (2013) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1301301110 · Cited by 392
Mega-scale experimental analysis of protein folding stability in biology and design
Tsuboyama, K., Dauparas, J., Chen, J., Laine, E., Mohseni Behbahani, Y., Weinstein, J. J., Mangan, N. M., Ovchinnikov, S., & Rocklin, G. J. (2023) · Nature · doi:10.1038/s41586-023-06328-6 · Cited by 350
Transposon-associated TnpB is a programmable RNA-guided DNA endonuclease
Karvelis, T., Druteika, G., Bigelyte, G., Budre, K., Zedaveinyte, R., Silanskas, A., Kazlauskas, D., Venclovas, Č., & Siksnys, V. (2021) · Nature · doi:10.1038/s41586-021-04058-1 · Cited by 348
GA4GH: International policies and standards for data sharing across genomic research and healthcare
Rehm, H. L., Page, A. J. H., Smith, L., Adams, J. B., Alterovitz, G., Babb, L. J., Barkley, M. P., Baudis, M., Beauvais, M. J. S., Beck, T., et al. (2021) · Cell Genomics · doi:10.1016/j.xgen.2021.100029 · Cited by 342
A Comprehensive, High-Resolution Map of a Gene’s Fitness Landscape
Firnberg, E., Labonte, J. W., Gray, J. J., & Ostermeier, M. (2014) · Molecular Biology and Evolution · doi:10.1093/molbev/msu081 · Cited by 340
CRISPRCasTyper: Automated Identification, Annotation, and Classification of CRISPR-Cas Loci
Russel, J., Pinilla‐Redondo, R., Mayo-Muñoz, D., Shah, S. A., & Sørensen, S. J. (2020) · The CRISPR Journal · doi:10.1089/crispr.2020.0059 · Cited by 312
GLU Variants Improve Transformer
Shazeer, N. (2020) · arXiv · doi:10.48550/arxiv.2002.05202 · Cited by 303
Direct Preference Optimization: Your Language Model is Secretly a Reward Model
Rafailov, R., Sharma, A., Mitchell, E., Ermon, S., Manning, C. D., & Finn, C. (2023) · arXiv · doi:10.48550/arxiv.2305.18290 · Cited by 293
Self-citationKotler, E., Shani, O., Goldfeld, G., Lotan‐Pompan, M., Tarcic, O., Gershoni, A., Hopf, T. A., Marks, D. S., Oren, M., & Segal, E. (2018) · Molecular Cell · doi:10.1016/j.molcel.2018.06.012 · Cited by 292
Evolutionary Implications of Microbial Genome Tetranucleotide Frequency Biases
Pride, D. T., Meinersmann, R. J., Wassenaar, T. M., & Blaser, M. J. (2003) · Genome Research · doi:10.1101/gr.335003 · Cited by 289
R2R - software to speed the depiction of aesthetic consensus RNA secondary structures
Weinberg, Z., & Breaker, R. R. (2011) · BMC Bioinformatics · doi:10.1186/1471-2105-12-3 · Cited by 287
Assessing the Impact of Secondary Structure and Solvent Accessibility on Protein Evolution
Goldman, N., Thorne, J. L., & Jones, D. T. (1998) · Genetics · doi:10.1093/genetics/149.1.445 · Cited by 281
Root Mean Square Layer Normalization
Zhang, B., & Sennrich, R. (2019) · arXiv · doi:10.48550/arxiv.1910.07467 · Cited by 262
Capturing the mutational landscape of the beta-lactamase TEM-1
Jacquier, H., Birgy, A., Nagard, H. L., Méchulam, Y., Schmitt, E., Glodt, J., Berçot, B., Petit, E., Poulain, J., Barnaud, G., et al. (2013) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1215206110 · Cited by 259
Chen, J., Hu, Z., Sun, S., Tan, Q., Wang, Y., Yu, Q., Zong, L., Hong, L., Xiao, J., Shen, T., et al. (2022) · doi:10.1101/2022.08.06.503062 · Cited by 247
Durrant, M. G., Li, M. M., Siranosian, B. A., Montgomery, S. B., & Bhatt, A. S. (2019) · Cell Host & Microbe · doi:10.1016/j.chom.2019.10.022 · Cited by 239
Self-citationCryptic genetic variation promotes rapid evolutionary adaptation in an RNA enzyme
Hayden, E. J., Ferrada, E., & Wagner, A. (2011) · Nature · doi:10.1038/nature10083 · Cited by 228
(2023) · doi:10.1101/2023.12.07.570727 · Cited by 207
LaFleur, T. L., Hossain, A., & Salis, H. M. (2022) · Nature Communications · doi:10.1038/s41467-022-32829-5 · Cited by 202
Diagonal State Spaces Are as Effective as Structured State Spaces
Gupta, A., Gu, A., & Berant, J. (2022) · neural information processing systems · doi:10.52202/068431-1670 · Cited by 192
Structural biology of CRISPR–Cas immunity and genome editing enzymes
Wang, J. Y., Pausch, P., & Doudna, J. A. (2022) · Nature Reviews Microbiology · doi:10.1038/s41579-022-00739-4 · Cited by 191
On the Parameterization and Initialization of Diagonal State Space Models
Gu, A., Goel, K., Gupta, A., & Ré, C. (2022) · neural information processing systems · doi:10.52202/068431-2607 · Cited by 183
Self-citationA catalogue of biochemically diverse CRISPR-Cas9 orthologs
Gasiūnas, G., Young, J. K., Karvelis, T., Kazlauskas, D., Urbaitis, T., Jasnauskaitė, M., Grusyte, M. M., Paulraj, S., Wang, P.-H., Hou, Z., et al. (2020) · Nature Communications · doi:10.1038/s41467-020-19344-1 · Cited by 181
DNABERT-2: Efficient Foundation Model and Benchmark For Multi-Species Genome
Zhou, Z., Ji, Y., Li, W., Dutta, P., Davuluri, R., & Liu, H. (2023) · arXiv · doi:10.48550/arxiv.2306.15006 · Cited by 179
Comprehensive mutational scanning of a kinase in vivo reveals substrate-dependent fitness landscapes
Melnikov, A., Rogov, P., Wang, L., Gnirke, A., & Mikkelsen, T. S. (2014) · Nucleic Acids Research · doi:10.1093/nar/gku511 · Cited by 177
Unsupervised evolution of protein and antibody complexes with a structure-informed language model
Shanker, V. R., Bruun, T. U. J., Hie, B., & Kim, P. S. (2024) · Science · doi:10.1126/science.adk8946 · Cited by 174
Self-citationHyenaDNA: Long-Range Genomic Sequence Modeling at Single Nucleotide Resolution
Nguyen, E., Poli, M., Faizi, M., Thomas, A., Birch-Sykes, C., Wornow, M., Patel, A., Rabideau, C., Massaroli, S., Bengio, Y., et al. (2023) · arXiv · doi:10.48550/arxiv.2306.15794 · Cited by 168
Self-citationGENA-LM: a family of open-source foundational DNA language models for long sequences
Fishman, V., Kuratov, Y., Shmelev, A., Petrov, M., Penzar, D., Shepelin, D., Chekanov, N., Kardymon, O. L., & Burtsev, M. (2024) · Nucleic Acids Research · doi:10.1093/nar/gkae1310 · Cited by 168
Gene essentiality determines chromosome organisation in bacteria
Rocha, E. P. C. (2003) · Nucleic Acids Research · doi:10.1093/nar/gkg859 · Cited by 166
An Automated Model Test System for Systematic Development and Improvement of Gene Expression Models
Reis, A. C., & Salis, H. M. (2020) · ACS Synthetic Biology · doi:10.1021/acssynbio.0c00394 · Cited by 166
Mega: Moving Average Equipped Gated AttentionPreprint
Ma, X., Zhou, C., Kong, X., He, J., Gui, L., Neubig, G., May, J., & Zettlemoyer, L. (2022) · arXiv (Cornell University) · doi:10.48550/arxiv.2209.10655 · Cited by 166
Bridge RNAs direct programmable recombination of target and donor DNA
Durrant, M. G., Perry, N. T., Pai, J. J., Jangid, A. R., Athukoralage, J. S., Hiraizumi, M., McSpedon, J. P., Pawluk, A., Nishimasu, H., Konermann, S., et al. (2024) · Nature · doi:10.1038/s41586-024-07552-4 · Cited by 163
Self-citationRapid Construction of Empirical RNA Fitness Landscapes
Pitt, J. N., & Ferré-D’Amaré, A. R. (2010) · Science · doi:10.1126/science.1192001 · Cited by 159
A DNA language model based on multispecies alignment predicts the effects of genome-wide variants
Benegas, G., Albors, C., Aw, A. J., Ye, C., & Song, Y. S. (2025) · Nature Biotechnology · doi:10.1038/s41587-024-02511-w · Cited by 145
Barabas, O., Ronning, D. R., Guynet, C., Hickman, A. B., Ton-Hoang, B., Chandler, M., & Dyda, F. (2008) · Cell · doi:10.1016/j.cell.2007.12.029 · Cited by 143
Protein language models learn evolutionary statistics of interacting sequence motifs
Zhang, Z., Wayment-Steele, H. K., Brixi, G., Wang, H., Kern, D., & Ovchinnikov, S. (2024) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2406285121 · Cited by 140
Self-citationGENA-LM: A Family of Open-Source Foundational DNA Language Models for Long Sequences
Fishman, V., Kuratov, Y., Shmelev, A., Petrov, M., Penzar, D., Shepelin, D., Chekanov, N., Kardymon, O., & Burtsev, M. (2023) · doi:10.1101/2023.06.12.544594 · Cited by 137 · cited 1×
“As a result, Transformer-based DNA models are constrained to short context lengths and use schemes that aggregate nucleotides into the basic units of language models, called tokens, thereby sacrificing single-nucleotide resolution ( 15 , 16 , 21 – 23 ).”
Pairwise and higher-order genetic interactions during the evolution of a tRNA
Domingo, J., Diss, G., & Lehner, B. (2018) · Nature · doi:10.1038/s41586-018-0170-7 · Cited by 137
Convolutions are competitive with transformers for protein sequence pretraining
Yang, K. K., Fusi, N., & Lu, A. X. (2024) · Cell Systems · doi:10.1016/j.cels.2024.01.008 · Cited by 137
Automated design of thousands of nonrepetitive parts for engineering stable genetic systems
Hossain, A., Lopez, E., Halper, S. M., Cetnar, D. P., Reis, A. C., Strickland, D., Klavins, E., & Salis, H. M. (2020) · Nature Biotechnology · doi:10.1038/s41587-020-0584-2 · Cited by 133
Hungry Hungry Hippos: Towards Language Modeling with State Space Models
Fu, D. Y., Dao, T., Saab, K. K., Thomas, A. W., Rudra, A., & Ré, C. (2022) · arXiv · doi:10.48550/arxiv.2212.14052 · Cited by 123
Self-citationGenSLMs: Genome-scale language models reveal SARS-CoV-2 evolutionary dynamics
Zvyagin, M., Brace, A., Hippe, K., Deng, Y., Zhang, B., Bohorquez, C. O., Clyde, A., Kale, B., Perez-Rivera, D., Ma, H., et al. (2023) · The International Journal of High Performance Computing Applications · doi:10.1177/10943420231201154 · Cited by 120
GenSLMs: Genome-scale language models reveal SARS-CoV-2 evolutionary dynamics
Zvyagin, M., Brace, A., Hippe, K., Deng, Y., Zhang, B., Bohorquez, C. O., Clyde, A., Kale, B., Perez-Rivera, D., Ma, H., et al. (2022) · doi:10.1101/2022.10.10.511571 · Cited by 120
GPN-MSA: an alignment-based DNA language model for genome-wide variant effect prediction
Benegas, G., Albors, C., Aw, A. J., Ye, C., & Song, Y. S. (2023) · doi:10.1101/2023.10.10.561776 · Cited by 118 · cited 1×
“For example, language models specifically trained on large corpuses of protein sequences or nucleotide coding sequences have demonstrated an ability to predict mutational effects on protein function ( 41 – 43 ) without any task-specific fine-tuning or supervision.”
Notin, P., Dias, M., Frazer, J., Marchena-Hurtado, J., Gomez, A., Marks, D. S., & Gal, Y. (2022) · arXiv · doi:10.48550/arxiv.2205.13760 · Cited by 118
Updated benchmarking of variant effect predictors using deep mutational scanning
Livesey, B., & Marsh, J. A. (2023) · Molecular Systems Biology · doi:10.15252/msb.202211474 · Cited by 118
Language Model Behavior: A Comprehensive Survey
Chang, T. A., & Bergen, B. K. (2024) · Computational Linguistics · doi:10.1162/coli_a_00492 · Cited by 105
Youngblut, N. D., de la Cuesta‐Zuluaga, J., Reischer, G. H., Dauser, S., Schuster, N., Walzer, C., Stalder, G. L., Farnleitner, A. H., & Ley, R. E. (2020) · mSystems · doi:10.1128/msystems.01045-20 · Cited by 100
Genomic language model predicts protein co-regulation and function
Hwang, Y., Cornman, A. L., Kellogg, E. H., Ovchinnikov, S., & Girguis, P. R. (2024) · Nature Communications · doi:10.1038/s41467-024-46947-9 · Cited by 99 · cited 1×
“As a result, Transformer-based DNA models are constrained to short context lengths and use schemes that aggregate nucleotides into the basic units of language models, called tokens, thereby sacrificing single-nucleotide resolution ( 15 , 16 , 21 – 23 ).”
Hyena Hierarchy: Towards Larger Convolutional Language Models
Poli, M., Massaroli, S., Nguyen, E., Fu, D. Y., Dao, T., Baccus, S., Bengio, Y., Ermon, S., & Ré, C. (2023) · arXiv · doi:10.48550/arxiv.2302.10866 · Cited by 96
Self-citationUrtecho, G., Tripp, A. D., Insigne, K. D., Kim, H., & Kosuri, S. (2018) · Biochemistry · doi:10.1021/acs.biochem.7b01069 · Cited by 92
Charformer: Fast Character Transformers via Gradient-based Subword Tokenization
Tay, Y., Tran, V. Q., Ruder, S., Gupta, J., Chung, H. W., Bahri, D., Qin, Z., Baumgartner, S., Yu, C., & Metzler, D. (2021) · arXiv · doi:10.48550/arxiv.2106.12672 · Cited by 92
Rockah-Shmuel, L., Tóth-Petróczy, Á., & Tawfik, D. S. (2015) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1004421 · Cited by 91
Codon language embeddings provide strong signals for use in protein engineering
Outeiral, C., & Deane, C. M. (2024) · Nature Machine Intelligence · doi:10.1038/s42256-024-00791-0 · Cited by 90
Effectively Modeling Time Series with Simple Discrete State SpacesPreprint
Zhang, M. S., Saab, K. K., Poli, M., Dao, T., Goel, K., & Ré, C. (2023) · International Conference on Learning Representations · doi:10.48550/arxiv.2303.09489 · Cited by 90
Self-citationProtein Model Discrimination Using Mutational Sensitivity Derived from Deep Sequencing
Adkar, B. V., Tripathi, A., Sahoo, A., Pahuja, K. B., Goswami, D., Chakrabarti, P., Swarnkar, M. K., Gokhale, R. S., & Varadarajan, R. (2012) · Structure · doi:10.1016/j.str.2011.11.021 · Cited by 87
Korkmaz, G., Holm, M., Wiens, T., & Sanyal, S. (2014) · Journal of Biological Chemistry · doi:10.1074/jbc.m114.606632 · Cited by 87
IMG/PR: a database of plasmids from genomes and metagenomes with rich annotations and metadata
Camargo, A. P., Call, L., Roux, S., Nayfach, S., Huntemann, M., Palaniappan, K., Ratner, A., Chu, K., Mukherjeep, S., Reddy, T. B. K., et al. (2024) · Nucleic Acids Research · doi:10.1093/nar/gkad964 · Cited by 87
Structure of PDE3A-SLFN12 complex reveals requirements for activation of SLFN12 RNase
Garvie, C. W., Wu, X., Papanastasiou, M., Lee, S., Fuller, J., Schnitzler, G. R., Horner, S. W., Baker, A., Zhang, T., Mullahoo, J. P., et al. (2021) · Nature Communications · doi:10.1038/s41467-021-24495-w · Cited by 85
Diversity, evolution, and classification of the RNA-guided nucleases TnpB and Cas12
Altae-Tran, H., Shmakov, S., Makarova, K. S., Wolf, Y. I., Kannan, S., Zhang, F., & Koonin, E. V. (2023) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2308224120 · Cited by 80
Guiding principles for the responsible development of artificial intelligence tools for healthcare
Badal, K., Lee, C. M., & Esserman, L. J. (2023) · Communications Medicine · doi:10.1038/s43856-023-00279-9 · Cited by 77
Guy, M. P., Young, D. L., Payea, M. J., Zhang, X., Kon, Y., Dean, K., Grayhack, E. J., Mathews, D. H., Fields, S., & Phizicky, E. M. (2014) · Genes & Development · doi:10.1101/gad.245936.114 · Cited by 76
Resurrecting Recurrent Neural Networks for Long Sequences
Orvieto, A., Smith, S. L., Gu, A., Fernando, A., Gulcehre, C., Pascanu, R., & De, S. (2023) · arXiv · doi:10.48550/arxiv.2303.06349 · Cited by 76
A proactive genotype-to-patient-phenotype map for cystathionine beta-synthase
Sun, S., Weile, J., Verby, M., Wu, Y., Wang, Y., Coté, A. G., Fotiadou, I., Kitaygorodsky, J., Vidal, M., Rine, J., et al. (2020) · Genome Medicine · doi:10.1186/s13073-020-0711-1 · Cited by 74
Direct and indirect impacts of synthetic biology on biodiversity conservation
Macfarlane, N. B. W., Adams, J. R., Bennett, E. L., Brooks, T. M., Delborne, J., Eggermont, H., Endy, D., Esvelt, K. M., Kolodziejczyk, B., Kuiken, T., et al. (2022) · iScience · doi:10.1016/j.isci.2022.105423 · Cited by 73
Deep learning and CRISPR-Cas13d ortholog discovery for optimized RNA targeting
Wei, J., Lotfy, P., Faizi, K., Baungaard, S., Gibson, E., Wang, E., Slabodkin, H., Kinnaman, E., Chandrasekaran, S. S., Kitano, H. C., et al. (2023) · Cell Systems · doi:10.1016/j.cels.2023.11.006 · Cited by 71
Self-citationRNA Structural Determinants of Optimal Codons Revealed by MAGE-Seq
Kelsic, E. D., Chung, H., Cohen, N., Park, J., Wang, H. H., & Kishony, R. (2016) · Cell Systems · doi:10.1016/j.cels.2016.11.004 · Cited by 70
Deep learning and CRISPR-Cas13d ortholog discovery for optimized RNA targeting
Wei, J., Lotfy, P., Faizi, K., Baungaard, S., Gibson, E., Wang, E., Slabodkin, H., Kinnaman, E., Chandrasekaran, S., Kitano, H., et al. (2021) · doi:10.1101/2021.09.14.460134 · Cited by 67 · cited 2×
“Custom Cas13 pHMMs that were previously generated by our group were also used ( 101 ).”
Self-citationMorin, S., Segafredo, G., Piccolis, M., Das, A., Das, M., Loffredi, N., Larbi, A., Mwamelo, K., Villanueva, E., Nobre, S., et al. (2022) · The Lancet Global Health · doi:10.1016/s2214-109x(22)00460-0 · Cited by 67
CodonBERT large language model for mRNA vaccinesPreprint
Li, S., Moayedpour, S., Li, R., Bailey, M., Riahi, S., Kogler-Anele, L., Miladi, M., Miner, J. C., Pertuy, F., Zheng, D., et al. (2024) · Genome Research · doi:10.1101/gr.278870.123 · Cited by 66
HyenaDNA: Long-Range Genomic Sequence Modeling at Single Nucleotide Resolution
Nguyen, E., Poli, M., Faizi, M., Thomas, A., Wornow, M., Birch-Sykes, C., Massaroli, S., Patel, A., Rabideau, C., Bengio, Y., et al. (2023) · Advances in Neural Information Processing Systems 36 · doi:10.52202/075280-1872 · Cited by 66
Self-citationTransposon-encoded nucleases use guide RNAs to promote their selfish spread
Meers, C., Le, H. C., Pesari, S. R., Hoffmann, F. T., Walker, M. W. G., Gezelle, J., Tang, S., & Sternberg, S. H. (2023) · Nature · doi:10.1038/s41586-023-06597-1 · Cited by 60 · cited 2×
“4A) ( 66 – 69 ).”
High-throughput assay and engineering of self-cleaving ribozymes by sequencing
Kobori, S., Nomura, Y., Miu, A., & Yokobayashi, Y. (2015) · Nucleic Acids Research · doi:10.1093/nar/gkv265 · Cited by 59
Yu, T. C., Liu, W. L., Brinck, M. S., Davis, J. E., Shek, J., Bower, G., Einav, T., Insigne, K. D., Phillips, R., Kosuri, S., et al. (2021) · Nature Communications · doi:10.1038/s41467-020-20094-3 · Cited by 59
Protein design meets biosecurity
Baker, D. A., & Church, G. M. (2024) · Science · doi:10.1126/science.ado1671 · Cited by 56
Feature Reuse and Scaling: Understanding Transfer Learning with Protein Language Models
Li, F.-Z., Amini, A. P., Yue, Y., Yang, K. K., & Lu, A. X. (2024) · doi:10.1101/2024.02.05.578959 · Cited by 51
Extending Context Window of Large Language Models via Positional Interpolation
Chen, S., Wong, S., Chen, L., & Tian, Y. (2023) · arXiv · doi:10.48550/arxiv.2306.15595 · Cited by 50
Chen, J. Z., Fowler, D. M., & Tokuriki, N. (2020) · eLife · doi:10.7554/elife.56707 · Cited by 47
Biotite: new tools for a versatile Python bioinformatics library
Kunzmann, P., Müller, T. D., Greil, M., Krumbach, J. H., Anter, J. M., Bauer, D., Islam, F., & Hamacher, K. (2023) · BMC Bioinformatics · doi:10.1186/s12859-023-05345-6 · Cited by 45
Evolving Better RNAfold Structure Prediction
Langdon, W. B., Petke, J., & Lorenz, R. (2018) · Lecture notes in computer science · doi:10.1007/978-3-319-77553-1_14 · Cited by 42
CKConv: Continuous Kernel Convolution For Sequential Data
Romero, D. W., Kuzina, A., Bekkers, E. J., Tomczak, J. M., & Hoogendoorn, M. (2021) · arXiv · doi:10.48550/arxiv.2102.02611 · Cited by 39
Comprehensive sequence-to-function mapping of cofactor-dependent RNA catalysis in the glmS ribozyme
Andreasson, J. O. L., Savinov, A., Block, S. M., & Greenleaf, W. J. (2020) · Nature Communications · doi:10.1038/s41467-020-15540-1 · Cited by 38
Pilault, J., Fathi, M., Firat, O., Pal, C., Bacon, P.-L., & Goroshin, R. (2023) · Neural Information Processing Systems · doi:10.52202/075280-0319 · Cited by 34
Effective Long-Context Scaling of Foundation Models
Xiong, W., Liu, J., Molybog, I., Zhang, H., Bhargava, P., Hou, R., Martin, L., Rungta, R., Sankararaman, K. A., Oguz, B., et al. (2024) · Proceedings of the 2024 Conference of the North American Chapter of the Association for Computational Linguistics: Human Language Technologies (Volume 1: Long Papers) · doi:10.18653/v1/2024.naacl-long.260 · Cited by 33
DaSilva, L. F., Senan, S., Patel, Z. M., Reddy, A. J., Gabbita, S., Nussbaum, Z., Córdova, C. M. V., Wenteler, A., Weber, N., Tunjic, T. M., et al. (2024) · doi:10.1101/2024.02.01.578352 · Cited by 32
Monarch Mixer: A Simple Sub-Quadratic GEMM-Based Architecture
Fu, D. Y., Arora, S., Grogan, J., Johnson, I., Eyuboglu, E. S., Thomas, A. W., Spector, B., Poli, M., Rudra, A., & Ré, C. (2023) · neural information processing systems · doi:10.52202/075280-3390 · Cited by 32
Self-citationSystematic and scalable genome-wide essentiality mapping to identify nonessential genes in phages
Piya, D., Nolan, N., Moore, M. L., Ramirez Hernandez, L. A., Cress, B. F., Young, R., Arkin, A. P., & Mutalik, V. K. (2023) · PLOS Biology · doi:10.1371/journal.pbio.3002416 · Cited by 31
Ring Attention with Blockwise Transformers for Near-Infinite Context
Liu, H., Zaharia, M., & Abbeel, P. (2023) · arXiv · doi:10.48550/arxiv.2310.01889 · Cited by 23
The evolution of Mycoplasma genitalium
Blanchard, A., & Bébéar, C. (2011) · Annals of the New York Academy of Sciences · doi:10.1111/j.1749-6632.2011.06418.x · Cited by 22
CRISPRtracrRNA: robust approach for CRISPR tracrRNA detection
Mitrofanov, A., Ziemann, M., Alkhnbashi, O. S., Hess, W. R., & Backofen, R. (2022) · Bioinformatics · doi:10.1093/bioinformatics/btac466 · Cited by 16
Bacterial phylogenetic tree construction based on genomic translation stop signals
Xu, L., Kuo, J., Liu, J.-K., & Wong, T.-Y. (2012) · Microbial Informatics and Experimentation · doi:10.1186/2042-5783-2-6 · Cited by 13
Fitness and Functional Landscapes of theE. coliRNase III Genernc
Weeks, R., & Ostermeier, M. (2023) · Molecular Biology and Evolution · doi:10.1093/molbev/msad047 · Cited by 12
The carbon footprint of computational research
(2023) · Nature Computational Science · doi:10.1038/s43588-023-00506-2 · Cited by 11
Effective Long-Context Scaling of Foundation Models
Xiong, W., Liu, J., Molybog, I., Zhang, H., Bhargava, P., Hou, R., Martin, L., Rungta, R., Sankararaman, K. A., Oguz, B., et al. (2023) · arXiv · doi:10.48550/arxiv.2309.16039 · Cited by 9
Silverstein, R. A., Sun, S., Verby, M., Weile, J., Wu, Y., Gebbia, M., Fotiadou, I., Kitaygorodsky, J., & Roth, F. P. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.10.06.463360 · Cited by 8
Bridge RNAs direct modular and programmable recombination of target and donor DNAPreprint
Durrant, M. G., Perry, N. T., Pai, J. J., Jangid, A. R., Athukoralage, J. S., Hiraizumi, M., McSpedon, J. P., Pawluk, A., Nishimasu, H., Konermann, S., et al. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.01.24.577089 · Cited by 7
Self-citationRapid in vivo exploration of a 5S rRNA neutral network
Zhang, Z. D., Nayar, M., Ammons, D., Rampersad, J., & Fox, G. E. (2009) · Journal of Microbiological Methods · doi:10.1016/j.mimet.2008.10.010 · Cited by 6
regLM: Designing realistic regulatory DNA with autoregressive language models
Lal, A., Garfield, D., Biancalani, T., & Eraslan, G. (2024) · doi:10.1101/2024.02.14.580373 · Cited by 6
By any means necessary: why lowering insulin prices is relevant to racial health equity
Peek, M. E. (2021) · The Lancet · doi:10.1016/s0140-6736(21)02315-1 · Cited by 5
Laughing Hyena Distillery: Extracting Compact Recurrences From Convolutions
Massaroli, S., Poli, M., Fu, D., Kumbong, H., Parnichkun, R., Romero, D., Timalsina, A., McIntyre, Q., Chen, B., Rudra, A., et al. (2023) · Advances in Neural Information Processing Systems 36 · doi:10.52202/075280-0747 · Cited by 4
Self-citationZoology: Measuring and Improving Recall in Efficient Language Models
Arora, S., Eyuboglu, S., Timalsina, A., Johnson, I., Poli, M., Zou, J., Rudra, A., & Ré, C. (2023) · arXiv · doi:10.48550/arxiv.2312.04927 · Cited by 4
Self-citationLanguage Model Behavior: A Comprehensive SurveyPreprint
Chang, T. A., & Bergen, B. K. (2023) · arXiv (Cornell University) · doi:10.48550/arxiv.2303.11504 · Cited by 3
Understanding In-Context Learning in Transformers and LLMs by Learning to Learn Discrete Functions
Bhattamishra, S., Patel, A., Blunsom, P., & Kanade, V. (2023) · arXiv · doi:10.48550/arxiv.2310.03016 · Cited by 3
Code for paper "Sequence modeling and design from molecular to genome scale with Evo"
Hie, B. (2024) · Zenodo · doi:10.5281/zenodo.12693561
Self-citation
Research works printed but not linked: 1
- N. L. Craig, M. Chandler, M. Gellert, A. M. Lambowitz, P. A. Rice, S. B. Sandmeyer, Eds., Mobile DNA III (Wiley, ed. 3, 2020).
Cited by
Sorted by citations
Genome modelling and design across all domains of life with Evo 2
Brixi, G., Durrant, M. G., Ku, J., Naghipourfar, M., Poli, M., Sun, G., Brockman, G., Chang, D., Fanton, A., Gonzalez, G. A., et al. (2026) · Nature · doi:10.1038/s41586-026-10176-5 · Cited by 544
Self-citationHow to build the virtual cell with artificial intelligence: Priorities and opportunities
Bunne, C., Roohani, Y., Rosen, Y., Gupta, A., Zhang, X., Roed, M., Alexandrov, T., AlQuraishi, M., Brennan, P., Burkhardt, D. B., et al. (2024) · Cell · doi:10.1016/j.cell.2024.11.015 · Cited by 393
Self-citationDesign of highly functional genome editors by modelling CRISPR–Cas sequences
Ruffolo, J. A., Nayfach, S., Gallagher, J. P., Bhatnagar, A., Beazer, J., Hussain, R., Russ, J., Yip, J., Hill, E., Pačesa, M., et al. (2025) · Nature · doi:10.1038/s41586-025-09298-z · Cited by 233
Scientific Large Language Models: A Survey on Biological & Chemical Domains
Zhang, Q., Ding, K., Lv, T., Wang, X., Yin, Q., Zhang, Y., Jing, Y., Wang, Y., Li, X., Xiang, Z., et al. (2025) · ACM Computing Surveys · doi:10.1145/3715318 · Cited by 216
Transformers and genome language models
Consens, M. E., Dufault, C., Wainberg, M., Forster, D., Karimzadeh, M., Goodarzi, H., Theis, F. J., Moses, A. M., & Wang, B. (2025) · Nature Machine Intelligence · doi:10.1038/s42256-025-01007-9 · Cited by 170
Metadata
- Authors
- Nguyen, E., Poli, M., Durrant, M. G., Kang, B., Katrekar, D., Li, D. B., Bartie, L. J., Thomas, A. W., King, S. H., Brixi, G., Sullivan, J., Ng, M. Y., Lewis, A., Lou, A., Ermon, S., Baccus, S. A., Hernandez-Boussard, T., Ré, C., Hsu, P. D., & Hie, B. L.
- Affiliations
- Arc Research Institute
- Stanford University (US)
- Chan Zuckerberg Biohub San Francisco (US)
- Stanford Medicine (US)
- Department of Biomedical Data Science, Stanford University, Stanford, CA, USA.
- Corresponding author
- Nguyen, Eric
- Poli, Michael
- Durrant, Matthew G.
- Kang, Brian
- Katrekar, Dhruva
- Li, David B.
- Hsu, Patrick D.
- Hie, Brian L.
- ORCID
- Venue
- Science
- Published
- 2024-11-15
- Volume
- 386
- Issue
- 6723
- Pages
- eado9336-eado9336
- ISSN
- 0036-8075 (ISSN-L)
- 1095-9203
- PMC
- PMC12057570
- OpenAlex
- W4404349982
- Semantic Scholar
- ed4a545249106483bb2adb80287728a577f340c1
- Cited by
- 580
- References
- 161
- Citations by year
- 2026: 273 · 2025: 246 · 2024: 61
- FWCI
- 66.79
- Type
- journal article
- Language
- en
- Open access
- green
- doi.org
- Licence
- cc-by
- Locations
- Science (publishedVersion)
- bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by)
- PubMed (publishedVersion)
- PubMed Central (submittedVersion)
- Funders
- National Institutes of Health (UL1TR003142)
- National Institute of Biomedical Imaging and Bioengineering (U54 EB020405)
- National Center for Advancing Translational Sciences (UM1TR004921)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Genomics and Phylogenetic Studies
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Machine Learning in Bioinformatics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › CRISPR and Genetic Engineering
- Keywords
- genomic foundation models, long-context language models, prokaryotic genomes, phage genome, scaling laws, genome organization
- MeSH terms
- DNA, DNA Transposable Elements, Models, Genetic, Mutation, RNA, Mutagenesis, Genome, Viral, Genome, Bacterial, Genome, Archaeal, Genomics, High-Throughput Nucleotide Sequencing, CRISPR-Cas Systems
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.
A language model drew the claims from the paper on 2026-10-05, under version 3 of the claim definition (model deepseek-v4.1-flash, prompt extract-v3).
A second model labelled each claim on 2026-10-05 (model jev-1.13-20260917, definition revision gold-guideline-v0.2).
Cited sources are checked with a language model’s help (gpt-6-sol).
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 ›
Are you an author, or do you see a problem with this page? Request a review