A General Harness for Protein Foundation Model Fitness Prediction
Organizations: Shanghai Jiao Tong University · Shanghai Innovation Institute · Shanghai Artificial Intelligence Laboratory
Abstract
Accurate fitness prediction is central to protein engineering and understanding sequence-function relationships. With advances in deep learning, protein foundation models (PFMs) have become widely used for this task. Recent analyses, however, show that these models share preferences reflecting their training corpora, while unreliable inputs can further distort fitness predictions. Family-specific evolutionary evidence and structural context can help address these limitations by providing complementary constraints on model scores, motivating VenusREM-Harness (VRH), a general, model-agnostic, training-free Retrieval-Enhanced Mutation harness. It fuses frozen model scores with multiple sequence alignment (MSA) evidence according to model uncertainty, then applies gated background correction and score shrinkage based on structural confidence and solvent exposure. Across 1,211 assays and 3.1 million measured variants from ProteinGym, VenusMutHub, and the newly curated viral benchmark VenusViroHub, all 71 configurations improve Spearman correlation on all 3 benchmarks by 0.073 on average, with broad gains across 5 metrics. Extended analyses relate retrieval gains to model-MSA preference differences, assess domain-level gains and immune-escape cases, and quantify computational speedups. Built with VRH, VenusREM2 is the first to rank highest in all function, taxon, MSA-depth, and mutation-depth categories, with a ProteinGym Average Spearman of 0.556, 0.038 above the prior best.
Figures & tables
| Component | Setting | PG Validation (21) | PG Full (217) | VMH Full (905) | VVH (89) | ||||
|---|---|---|---|---|---|---|---|---|---|
| Score | Score | Score | Score | ||||||
| Raw | Native field | 0.5573 | — | 0.5069 | — | 0.2126 | — | 0.2709 | — |
| MSA fusion | w/o Ent. Weight. | 0.5742 | +0.017 | 0.5205 | +0.014 | 0.2341 | +0.022 | 0.2897 | +0.019 |
| w/ Ent. Weight. | 0.5774 | +0.020 | 0.5246 | +0.018 | 0.2336 | +0.021 | 0.2901 | +0.019 | |
| Model calibration | w/o Coh. Gating | 0.5641 | +0.007 | 0.5219 | +0.015 | 0.2326 | +0.020 | 0.2841 | +0.013 |
| w/ Coh. Gating | 0.5788 | +0.022 | 0.5284 | +0.022 | 0.2369 | +0.024 | 0.2942 | +0.023 | |
Appendix figures & tables68 assets
Supplementary material from the paper’s appendix.
Appendix
| Dataset | assays | Sequence length | MSA depth | MSA occupancy | Mean pLDDT | Mean RSA |
|---|---|---|---|---|---|---|
| ProteinGym | 217 | 397 502 | 8,650 6,713 | 0.67 0.23 | 86.6 10.8 | 0.37 0.09 |
| VenusMutHub | 905 | 288 239 | 9,791 6,452 | 0.69 0.18 | 92.1 7.5 | 0.32 0.08 |
| VenusViroHub | 89 | 559 383 | 1,217 2,610 | 0.62 0.14 | 83.9 9.8 | 0.33 0.07 |
| Dataset | Unique WT | Structures | pLDDT | 50–70 | 70–90 | |
|---|---|---|---|---|---|---|
| ProteinGym | 187 | 187 | 18.9% | 9.0% | 22.6% | 49.5% |
| VenusMutHub | 527 | 527 | 6.7% | 3.3% | 10.6% | 79.5% |
| VenusViroHub | 52 | 52 | 9.7% | 13.2% | 36.0% | 41.1% |
| Dataset | RSA | RSA – | RSA |
|---|---|---|---|
| ProteinGym | 36.7% | 29.4% | 33.9% |
| VenusMutHub | 45.8% | 28.9% | 25.3% |
| VenusViroHub | 40.6% | 30.2% | 29.2% |
| Harmonised phenotype | Assays | Evaluated substitutions |
|---|---|---|
| Immune escape | 21 | 148,468 |
| Cell entry | 18 | 189,048 |
| Binding | 17 | 90,245 |
| Expression | 15 | 58,120 |
| Fitness | 15 | 114,287 |
| Stability | 2 | 11,187 |
| Virus and construct | Source |
|---|---|
| HIV-1 BF520 Env | ( Radford & Bloom, 2025 ) |
| HIV-1 TRO11 Env | ( Radford & Bloom, 2025 ) |
| Influenza A/H3N2 HK19 HA | ( Welsh et al., 2024 ) |
| Influenza A/H3N2 MC22 HA | ( Yu et al., 2025 ) |
| Lassa virus Josiah GP | ( Carr et al., 2024 ) |
| Nipah virus F | ( Larsen et al., 2026 ) |
| Virus and construct | Source |
|---|---|
| HIV-1 BF520 Env | ( Radford et al., 2023 ) |
| Human coronavirus 229E spike | ( Harari et al., 2026 ) |
| Influenza A/H5N1 HA | ( Dadonaite et al., 2024a ) |
| Lassa virus Josiah GP | ( Carr & Bloom, 2026 ) † |
| Respiratory syncytial virus A F | ( Simonich et al., 2026 ) |
| SARS-CoV-2 XBB.1.5 RBD | ( Dadonaite et al., 2024b ) |
| Virus and construct | Source |
|---|---|
| Chikungunya virus E3/E2/E1 | ( Ju et al., 2025 ) |
| Chikungunya virus E3/E2/E1 | ( Ju et al., 2025 ) |
| HIV-1 BF520 Env | ( Radford & Bloom, 2025 ) |
| HIV-1 TRO11 Env | ( Radford & Bloom, 2025 ) |
| Human coronavirus 229E spike | ( Harari et al., 2026 ) |
| Influenza A/H3N2 MC22 HA | ( Yu et al., 2025 ) |
| Virus and construct | Source |
|---|---|
| Chikungunya virus E3/E2/E1 | ( Ju et al., 2025 ) |
| Human coronavirus 229E spike | ( Harari et al., 2026 ) |
| Nipah virus Malaysia RBP | ( Larsen et al., 2025 ) |
| SARS-CoV-2 Alpha RBD | ( Starr et al., 2022a ) |
| SARS-CoV-2 Beta RBD | ( Starr et al., 2022a ) |
| SARS-CoV-2 Delta RBD | ( Starr et al., 2022a ) |
| Virus and construct | Source |
|---|---|
| Bat coronavirus PRD0038 RBD | ( Starr et al., 2022c ) |
| Bat coronavirus RmYN02 RBD | ( Starr et al., 2022c ) |
| Bat coronavirus RsYN04 RBD | ( Starr et al., 2022c ) |
| SARS-CoV-2 PLpro | ( Wu et al., 2024 ) |
| SARS-CoV-2 Alpha RBD | ( Starr et al., 2022a ) |
| SARS-CoV-2 Beta RBD | ( Starr et al., 2022a ) |
| Virus and construct | Source |
|---|---|
| Coxsackievirus B3 2A | ( Álvarez-Rodríguez et al., 2024 ) |
| Coxsackievirus B3 2B | ( Álvarez-Rodríguez et al., 2024 ) |
| Coxsackievirus B3 2C | ( Álvarez-Rodríguez et al., 2024 ) |
| Coxsackievirus B3 3A | ( Álvarez-Rodríguez et al., 2024 ) |
| Coxsackievirus B3 3B | ( Álvarez-Rodríguez et al., 2024 ) |
| Coxsackievirus B3 3C | ( Álvarez-Rodríguez et al., 2024 ) |
| Virus and construct | Source |
|---|---|
| Influenza A/H3N2 MC22 HA | ( Yu et al., 2025 ) |
| Influenza A/H5N1 HA | ( Dadonaite et al., 2024a ) |
| SARS-CoV-2 PLpro | ( Wu et al., 2024 ) |
| Model | Family | Evaluated variants | Readout | Source |
|---|---|---|---|---|
| ESM-1b | Masked sequence LM | 650M | mask, wt | Official repository |
| ESM-1v | Masked sequence LM | 650M | mask, wt | Official repository |
| ESM-2 | Masked sequence LM | 8M/35M/150M/650M/3B | mask, wt | Official repository |
| ESMC | Masked sequence LM | 300M, 600M | mask | Official repository |
| CARP | Masked sequence LM | 600K/38M/76M/640M | wt | Official repository |
| S3F | Structure-aware | S3F | wt, mask | Official repository |
| Rank | Model | Seq | Str | Evo | Average | Activity | Binding | Expression | Organismal | Stability |
|---|---|---|---|---|---|---|---|---|---|---|
| — | VenusREM2 (ours) | ✓ | ✓ | ✓ | 0.556 | 0.541 | 0.495 | 0.557 | 0.494 | 0.691 |
| 1 | AIDO Protein-RAG (16B) ( Sun et al., 2024 ) | — | ✓ | ✓ | 0.518 | 0.517 | 0.426 | 0.522 | 0.491 | 0.635 |
| 2 | VenusREM ( Tan et al., 2025b ) | ✓ | ✓ | ✓ | 0.518 | 0.495 | 0.454 | 0.533 | 0.459 | 0.650 |
| 3 | ProSST ( ) ( Li et al., 2024 ) | ✓ | ✓ | — | 0.507 | 0.476 | 0.445 | 0.530 | 0.431 | 0.653 |
| 5 | S3F-MSA ( Zhang et al., 2024 ) | — | ✓ | ✓ | 0.496 | 0.502 | 0.440 | 0.479 | 0.477 | 0.581 |
| 8 | Protriever ( Weitzman et al., 2025 ) | — | — | ✓ | 0.479 | 0.487 | 0.396 | 0.496 | 0.479 | 0.537 |
| Rank | Model | Seq | Str | Evo | Average | Activity | Binding | Expression | Organismal | Stability |
|---|---|---|---|---|---|---|---|---|---|---|
| — | VenusREM2 (ours) | ✓ | ✓ | ✓ | 0.556 | 0.541 | 0.495 | 0.557 | 0.494 | 0.691 |
| 1 | AIDO Protein-RAG (16B) ( Sun et al., 2024 ) | — | ✓ | ✓ | 0.518 | 0.517 | 0.426 | 0.522 | 0.491 | 0.635 |
| 2 | VenusREM ( Tan et al., 2025b ) | ✓ | ✓ | ✓ | 0.518 | 0.495 | 0.454 | 0.533 | 0.459 | 0.650 |
| 3 | ProSST ( ) ( Li et al., 2024 ) | ✓ | ✓ | — | 0.507 | 0.476 | 0.445 | 0.530 | 0.431 | 0.653 |
| 5 | S3F-MSA ( Zhang et al., 2024 ) | — | ✓ | ✓ | 0.496 | 0.502 | 0.440 | 0.479 | 0.477 | 0.581 |
| 8 | Protriever ( Weitzman et al., 2025 ) | — | — | ✓ | 0.479 | 0.487 | 0.396 | 0.496 | 0.479 | 0.537 |
| Rank | Model | Seq | Str | Evo | Human | Other euk. | Prok. | Virus | Low | Medium | High |
|---|---|---|---|---|---|---|---|---|---|---|---|
| — | VenusREM2 (ours) | ✓ | ✓ | ✓ | 0.562 | 0.615 | 0.601 | 0.540 | 0.553 | 0.557 | 0.623 |
| 1 | AIDO Protein-RAG (16B) ( Sun et al., 2024 ) | — | ✓ | ✓ | 0.531 | 0.587 | 0.558 | 0.522 | 0.498 | 0.534 | 0.585 |
| 2 | VenusREM ( Tan et al., 2025b ) | ✓ | ✓ | ✓ | 0.529 | 0.582 | 0.549 | 0.492 | 0.495 | 0.524 | 0.577 |
| 3 | ProSST ( ) ( Li et al., 2024 ) | ✓ | ✓ | — | 0.516 | 0.573 | 0.549 | 0.454 | 0.465 | 0.507 | 0.580 |
| 5 | S3F-MSA ( Zhang et al., 2024 ) | — | ✓ | ✓ | 0.502 | 0.558 | 0.521 | 0.502 | 0.469 | 0.509 | 0.547 |
| 8 | Protriever ( Weitzman et al., 2025 ) | — | — | ✓ | 0.480 | 0.542 | 0.492 | 0.516 | 0.464 | 0.498 | 0.512 |
| Stratification | Group | Raw | VRH | |
|---|---|---|---|---|
| Taxon | Human | 0.524 | 0.562 | +0.038 |
| Other eukaryotes | 0.596 | 0.615 | +0.019 | |
| Prokaryote | 0.579 | 0.601 | +0.022 | |
| Virus | 0.476 | 0.540 | +0.064 | |
| MSA depth | Low | 0.498 | 0.553 | +0.055 |
| Medium | 0.517 | 0.557 | +0.040 |
| Rank | Model | Seq | Str | Evo | 1 | 2 | 3 | 4 | |
|---|---|---|---|---|---|---|---|---|---|
| — | VenusREM2 (ours) | ✓ | ✓ | ✓ | 0.576 | 0.473 | 0.468 | 0.424 | 0.473 |
| 1 | AIDO Protein-RAG (16B) ( Sun et al., 2024 ) | — | ✓ | ✓ | 0.527 | 0.414 | 0.419 | 0.394 | 0.414 |
| 2 | VenusREM ( Tan et al., 2025b ) | ✓ | ✓ | ✓ | 0.534 | 0.397 | 0.355 | 0.322 | 0.368 |
| 3 | ProSST ( ) ( Li et al., 2024 ) | ✓ | ✓ | — | 0.521 | 0.394 | 0.317 | 0.277 | 0.332 |
| 5 | S3F-MSA ( Zhang et al., 2024 ) | — | ✓ | ✓ | 0.499 | 0.333 | 0.378 | 0.346 | 0.383 |
| 8 | Protriever ( Weitzman et al., 2025 ) | — | — | ✓ | 0.473 | 0.320 | 0.427 | 0.387 | 0.427 |
| Dataset | Pairwise | Corpus | Native–MSA | Complete sites | Aligned sites | |
|---|---|---|---|---|---|---|
| Median | Pooled | (%) | ||||
| ProteinGym | 0.995 | 0.991 | 0.965 | 0.758 | 27,763 | 19.7 |
| VenusMutHub | 0.996 | 0.989 | 0.964 | 0.863 | 148,983 | 24.8 |
| VenusViroHub | 0.990 | 0.885 | 0.976 | 0.621 | 29,416 | 38.1 |
| Family | Raw | VRH | Mean change | Median change | Wins / | |
|---|---|---|---|---|---|---|
| Structure-aware | 26 | 0.450 | 0.485 | +0.035 | +0.029 | 26 / 26 |
| Masked sequence LM | 20 | 0.359 | 0.440 | +0.081 | +0.062 | 20 / 20 |
| Inverse folding | 10 | 0.382 | 0.447 | +0.065 | +0.070 | 10 / 10 |
| Autoregressive LM | 15 | 0.302 | 0.410 | +0.108 | +0.098 | 15 / 15 |
| Metric | Raw | VRH | |
|---|---|---|---|
| Spearman | 0.541 0.180 | 0.575 0.159 | +0.034 |
| NDCG | 0.794 0.138 | 0.813 0.130 | +0.019 |
| AUC | 0.796 0.100 | 0.815 0.091 | +0.019 |
| MCC | 0.422 0.164 | 0.450 0.151 | +0.028 |
| Top-recall | 0.261 0.131 | 0.270 0.131 | +0.008 |
| Category | Structure (26) | Masked LM (20) | Inverse folding (10) | AR LM (15) | All (71) | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| Activity | 0.442 | 0.486 | 0.371 | 0.453 | 0.318 | 0.414 | 0.321 | 0.434 | 0.379 | 0.456 |
| Binding | 0.381 | 0.407 | 0.288 | 0.353 | 0.327 | 0.384 | 0.251 | 0.313 | 0.320 | 0.369 |
| Expression | 0.459 | 0.487 | 0.369 | 0.433 | 0.382 | 0.429 | 0.334 | 0.400 | 0.397 | 0.445 |
| Organismal Fitness | 0.378 | 0.423 | 0.295 | 0.394 | 0.294 | 0.373 | 0.334 | 0.397 | 0.334 | 0.402 |
| Stability | 0.589 | 0.623 | 0.474 | 0.567 | 0.588 | 0.635 | 0.272 | 0.507 | 0.489 | 0.584 |
| Family | Raw | VRH | Mean change | Median change | Wins / | |
|---|---|---|---|---|---|---|
| Structure-aware | 26 | 0.172 | 0.214 | +0.042 | +0.032 | 26 / 26 |
| Masked sequence LM | 20 | 0.134 | 0.196 | +0.062 | +0.050 | 20 / 20 |
| Inverse folding | 10 | 0.210 | 0.256 | +0.046 | +0.049 | 10 / 10 |
| Autoregressive LM | 15 | 0.102 | 0.182 | +0.080 | +0.070 | 15 / 15 |
| Family | Raw | VRH | Mean change | Median change | Wins / | |
|---|---|---|---|---|---|---|
| Structure-aware | 26 | 0.215 | 0.280 | +0.065 | +0.063 | 26 / 26 |
| Masked sequence LM | 20 | 0.072 | 0.222 | +0.151 | +0.147 | 20 / 20 |
| Inverse folding | 10 | 0.271 | 0.307 | +0.036 | +0.036 | 10 / 10 |
| Autoregressive LM | 15 | 0.099 | 0.206 | +0.107 | +0.110 | 15 / 15 |
| Benchmark | Resampling unit | Units | Mean | Wins / |
|---|---|---|---|---|
| ProteinGym | UniProt ID | 186 | +0.068 | 71 / 71 |
| VenusMutHub | wild-type sequence | 527 | +0.056 | 71 / 71 |
| VenusViroHub | phenotype–backbone cell | 52 | +0.094 | 71 / 71 |
| Comparison | Mean | Median | Wins / | Ties |
|---|---|---|---|---|
| Gated no correction | +0.012 | +0.010 | 71 / 71 | 0 |
| Ungated no correction | +0.003 | +0.003 | 43 / 71 | 0 |
| Gated ungated | +0.010 | +0.009 | 71 / 71 | 0 |
| Interface | Context at site | WT residue visible | Field construction | Multi-mutant scoring |
|---|---|---|---|---|
| Masked marginal | No | masked inputs | Additive native field | |
| Wild-type field | Full native | Yes | 1 forward pass | Additive native field |
| Teacher forcing | Prefix | No (prefix only) | 1 forward pass | Additive unless rescored |
| Dataset | Masked | Wild-type | ||||
|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | |||
| ProteinGym | 0.389 | 0.468 | +0.079 | 0.395 | 0.458 | +0.062 |
| VenusMutHub | 0.157 | 0.218 | +0.061 | 0.142 | 0.187 | +0.045 |
| VenusViroHub | 0.082 | 0.239 | +0.156 | 0.076 | 0.263 | +0.188 |
| Family | Model | Full sequence | Native field | Wins | |
|---|---|---|---|---|---|
| Inverse folding | ESM-IF1 | 0.422 | 0.421 | 95/217 | |
| Autoregressive | ProGen2 -S | 0.336 | 0.288 | 47/217 | |
| Autoregressive | ProGen2 -M | 0.380 | 0.337 | 54/217 | |
| Autoregressive | ProGen2 -B | 0.378 | 0.330 | 58/217 | |
| Autoregressive | ProGen2 -L | 0.380 | 0.327 | 53/217 | |
| Autoregressive | ProGen2 -XL | 0.391 | 0.351 | 67/217 |
| Route | Context | Evaluations | Order | Multi-mutant score |
|---|---|---|---|---|
| Official random order | Mutant + structure | Per mutant | Yes | May be non-additive |
| Native TF | Native + structure | Per protein | No | Additive |
| Conditional TF | Mutant background + structure | Per background | No | Non-additive |
| Subset | Quartets | Single TF | Single official | Double TF | Double official | official | conditional TF |
|---|---|---|---|---|---|---|---|
| All | 535,917 | 0.306 | 0.105 | 0.284 | 0.165 | 0.005 | 0.012 |
| Non-floor | 400,118 | 0.312 | 0.091 | 0.292 | 0.152 | 0.012 | 0.022 |
| Rank | Model | Readout | VRH | Overall | Activity | Binding | Expression | Organismal | Stability |
|---|---|---|---|---|---|---|---|---|---|
| 1 | VenusREM2 | wt | ✓ | 0.556 | 0.541 | 0.495 | 0.557 | 0.494 | 0.691 |
| 2 | ProSST (K=4096) | wt | ✓ | 0.541 | 0.516 | 0.494 | 0.543 | 0.475 | 0.674 |
| 3 | ProSST (K=2048) | wt | ✓ | 0.534 | 0.528 | 0.460 | 0.541 | 0.469 | 0.673 |
| 4 | ProSST (K=1024) | wt | ✓ | 0.532 | 0.513 | 0.467 | 0.533 | 0.476 | 0.670 |
| 5 | ProSST (K=128) | wt | ✓ | 0.525 | 0.508 | 0.472 | 0.517 | 0.463 | 0.663 |
| 6 | VenusREM2 | wt | 0.524 | 0.480 | 0.479 | 0.538 | 0.444 | 0.681 |
| Rank | Model | Readout | VRH | Overall | stability | activity | PPI bind. | selectivity | DTI bind. |
|---|---|---|---|---|---|---|---|---|---|
| 1 | ProteinMPNN (v_48_020) | tf | ✓ | 0.271 | 0.399 | 0.108 | 0.024 | 0.012 | 0.198 |
| 2 | ESM-IF1 | tf | ✓ | 0.267 | 0.356 | 0.152 | 0.108 | 0.018 | 0.269 |
| 3 | ProteinMPNN-Soluble (v_48_020) | tf | ✓ | 0.265 | 0.396 | 0.106 | 0.036 | 0.000 | 0.093 |
| 4 | ProteinMPNN-Soluble (v_48_030) | tf | ✓ | 0.265 | 0.383 | 0.120 | 0.039 | 0.022 | 0.168 |
| 5 | ProteinMPNN (v_48_030) | tf | ✓ | 0.259 | 0.388 | 0.106 | 0.019 | -0.020 | 0.137 |
| 6 | VenusREM2 | wt | ✓ | 0.258 | 0.351 | 0.122 | 0.113 | -0.017 | 0.274 |
| Rank | Model | Readout | VRH | Overall | fitness | activity | expression | cell entry | binding | stability | immune esc. |
|---|---|---|---|---|---|---|---|---|---|---|---|
| 1 | SaProt (650M_PDB, mask) | mask | ✓ | 0.320 | 0.394 | 0.577 | 0.569 | 0.484 | 0.145 | 0.045 | 0.026 |
| 2 | S3F (wt) | wt | ✓ | 0.318 | 0.404 | 0.582 | 0.540 | 0.493 | 0.111 | 0.052 | 0.046 |
| 3 | ESM-IF1 | tf | ✓ | 0.318 | 0.359 | 0.603 | 0.566 | 0.439 | 0.173 | 0.060 | 0.026 |
| 4 | MIF-ST | tf | ✓ | 0.317 | 0.391 | 0.542 | 0.553 | 0.476 | 0.144 | 0.068 | 0.042 |
| 5 | ProSST (K=2048) | wt | ✓ | 0.310 | 0.366 | 0.570 | 0.561 | 0.469 | 0.126 | 0.048 | 0.032 |
| 6 | SaProt (650M_PDB, wt) | wt | ✓ | 0.307 | 0.343 | 0.608 | 0.564 | 0.432 | 0.131 | 0.052 | 0.022 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ESM-IF1 | 0.421 | 0.466 | 0.457 | 0.467 | 0.473 | 0.474 |
| MIF-ST | 0.430 | 0.440 | 0.441 | 0.444 | 0.456 | 0.458 |
| ProteinMPNN (v_48_002) | 0.378 | 0.425 | 0.417 | 0.431 | 0.441 | 0.443 |
| ProteinMPNN (v_48_010) | 0.385 | 0.431 | 0.424 | 0.439 | 0.449 | 0.451 |
| ProteinMPNN (v_48_020) | 0.389 | 0.434 | 0.430 | 0.444 | 0.454 | 0.456 |
| ProteinMPNN (v_48_030) | 0.381 | 0.428 | 0.431 | 0.440 | 0.451 | 0.453 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ESM3 | 0.442 | 0.461 | 0.460 | 0.469 | 0.481 | 0.483 |
| ProSST (K=1024) | 0.485 | 0.517 | 0.513 | 0.522 | 0.530 | 0.532 |
| ProSST (K=128) | 0.469 | 0.506 | 0.503 | 0.512 | 0.522 | 0.525 |
| ProSST (K=20) | 0.438 | 0.486 | 0.481 | 0.495 | 0.507 | 0.510 |
| ProSST (K=2048) | 0.507 | 0.525 | 0.522 | 0.528 | 0.533 | 0.534 |
| ProSST (K=4096) | 0.498 | 0.525 | 0.522 | 0.531 | 0.538 | 0.541 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ProGen2-B | 0.330 | 0.377 | 0.387 | 0.396 | 0.419 | 0.422 |
| ProGen2-L | 0.327 | 0.371 | 0.382 | 0.390 | 0.413 | 0.416 |
| ProGen2-M | 0.337 | 0.380 | 0.391 | 0.398 | 0.420 | 0.423 |
| ProGen2-S | 0.288 | 0.362 | 0.378 | 0.386 | 0.404 | 0.406 |
| ProGen2-XL | 0.351 | 0.373 | 0.375 | 0.385 | 0.415 | 0.419 |
| ProGen3-112M | 0.232 | 0.341 | 0.367 | 0.381 | 0.398 | 0.401 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| CARP-38M | 0.314 | 0.342 | 0.356 | 0.367 | 0.409 | 0.415 |
| CARP-600K | 0.159 | 0.221 | 0.249 | 0.271 | 0.345 | 0.354 |
| CARP-640M | 0.390 | 0.427 | 0.433 | 0.437 | 0.450 | 0.451 |
| CARP-76M | 0.360 | 0.380 | 0.391 | 0.399 | 0.432 | 0.436 |
| ESM-1b (mask) | 0.389 | 0.430 | 0.436 | 0.440 | 0.453 | 0.455 |
| ESM-1b (wt) | 0.394 | 0.407 | 0.414 | 0.418 | 0.445 | 0.447 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ESM-IF1 | 0.216 | 0.260 | 0.243 | 0.260 | 0.268 | 0.267 |
| MIF-ST | 0.196 | 0.209 | 0.203 | 0.210 | 0.228 | 0.230 |
| ProteinMPNN (v_48_002) | 0.191 | 0.233 | 0.232 | 0.241 | 0.245 | 0.244 |
| ProteinMPNN (v_48_010) | 0.218 | 0.240 | 0.246 | 0.247 | 0.254 | 0.256 |
| ProteinMPNN (v_48_020) | 0.221 | 0.257 | 0.254 | 0.262 | 0.269 | 0.271 |
| ProteinMPNN (v_48_030) | 0.210 | 0.244 | 0.251 | 0.251 | 0.260 | 0.259 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ESM3 | 0.185 | 0.210 | 0.214 | 0.213 | 0.226 | 0.226 |
| ProSST (K=1024) | 0.162 | 0.223 | 0.236 | 0.231 | 0.241 | 0.242 |
| ProSST (K=128) | 0.139 | 0.213 | 0.225 | 0.221 | 0.231 | 0.230 |
| ProSST (K=20) | 0.143 | 0.213 | 0.216 | 0.224 | 0.237 | 0.238 |
| ProSST (K=2048) | 0.213 | 0.234 | 0.233 | 0.237 | 0.250 | 0.249 |
| ProSST (K=4096) | 0.161 | 0.225 | 0.228 | 0.230 | 0.238 | 0.237 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ProGen2-B | 0.124 | 0.168 | 0.183 | 0.180 | 0.193 | 0.194 |
| ProGen2-L | 0.120 | 0.159 | 0.175 | 0.175 | 0.189 | 0.188 |
| ProGen2-M | 0.115 | 0.154 | 0.173 | 0.167 | 0.182 | 0.182 |
| ProGen2-S | 0.091 | 0.149 | 0.179 | 0.179 | 0.191 | 0.192 |
| ProGen2-XL | 0.123 | 0.148 | 0.149 | 0.153 | 0.178 | 0.176 |
| ProGen3-112M | 0.072 | 0.141 | 0.160 | 0.162 | 0.176 | 0.177 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| CARP-38M | 0.073 | 0.102 | 0.136 | 0.132 | 0.159 | 0.158 |
| CARP-600K | 0.021 | 0.063 | 0.120 | 0.115 | 0.150 | 0.151 |
| CARP-640M | 0.157 | 0.187 | 0.191 | 0.194 | 0.206 | 0.206 |
| CARP-76M | 0.131 | 0.149 | 0.172 | 0.174 | 0.192 | 0.194 |
| ESM-1b (mask) | 0.166 | 0.198 | 0.200 | 0.200 | 0.214 | 0.211 |
| ESM-1b (wt) | 0.153 | 0.166 | 0.174 | 0.171 | 0.194 | 0.193 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ESM-IF1 | 0.285 | 0.305 | 0.300 | 0.307 | 0.316 | 0.318 |
| MIF-ST | 0.288 | 0.299 | 0.304 | 0.303 | 0.316 | 0.317 |
| ProteinMPNN (v_48_002) | 0.269 | 0.285 | 0.283 | 0.291 | 0.304 | 0.305 |
| ProteinMPNN (v_48_010) | 0.271 | 0.285 | 0.285 | 0.292 | 0.305 | 0.306 |
| ProteinMPNN (v_48_020) | 0.270 | 0.284 | 0.285 | 0.291 | 0.304 | 0.306 |
| ProteinMPNN (v_48_030) | 0.266 | 0.280 | 0.284 | 0.288 | 0.302 | 0.304 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ESM3 | 0.137 | 0.183 | 0.196 | 0.211 | 0.244 | 0.249 |
| ProSST (K=1024) | 0.204 | 0.243 | 0.247 | 0.250 | 0.272 | 0.275 |
| ProSST (K=128) | 0.179 | 0.223 | 0.232 | 0.234 | 0.261 | 0.264 |
| ProSST (K=20) | 0.161 | 0.211 | 0.213 | 0.224 | 0.254 | 0.259 |
| ProSST (K=2048) | 0.271 | 0.290 | 0.284 | 0.294 | 0.309 | 0.310 |
| ProSST (K=4096) | 0.200 | 0.237 | 0.242 | 0.244 | 0.268 | 0.271 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| ProGen2-B | 0.076 | 0.123 | 0.132 | 0.136 | 0.172 | 0.175 |
| ProGen2-L | 0.064 | 0.121 | 0.136 | 0.137 | 0.169 | 0.173 |
| ProGen2-M | 0.117 | 0.152 | 0.159 | 0.165 | 0.208 | 0.212 |
| ProGen2-S | 0.045 | 0.114 | 0.128 | 0.133 | 0.167 | 0.171 |
| ProGen2-XL | 0.162 | 0.179 | 0.179 | 0.186 | 0.232 | 0.235 |
| ProGen3-112M | 0.034 | 0.117 | 0.152 | 0.159 | 0.178 | 0.182 |
| Model | Raw | Adaptive mix | + Ungated CCD | + Gated CCD | + RSA | + pLDDT (Full) |
|---|---|---|---|---|---|---|
| CARP-38M | 0.027 | 0.067 | 0.096 | 0.112 | 0.213 | 0.222 |
| CARP-600K | 0.006 | 0.047 | 0.087 | 0.098 | 0.201 | 0.210 |
| CARP-640M | 0.092 | 0.162 | 0.185 | 0.190 | 0.212 | 0.215 |
| CARP-76M | 0.050 | 0.090 | 0.113 | 0.130 | 0.221 | 0.230 |
| ESM-1b (mask) | 0.093 | 0.169 | 0.192 | 0.197 | 0.220 | 0.223 |
| ESM-1b (wt) | 0.094 | 0.127 | 0.150 | 0.159 | 0.240 | 0.248 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ESM3 | 0.442 | 0.483 | 0.767 | 0.780 | 0.744 | 0.765 | 0.346 | 0.377 | 0.228 | 0.236 |
| ProSST (K=1024) | 0.485 | 0.532 | 0.761 | 0.790 | 0.764 | 0.788 | 0.372 | 0.410 | 0.230 | 0.251 |
| ProSST (K=128) | 0.469 | 0.525 | 0.754 | 0.789 | 0.757 | 0.786 | 0.363 | 0.407 | 0.227 | 0.253 |
| ProSST (K=20) | 0.438 | 0.510 | 0.745 | 0.786 | 0.739 | 0.777 | 0.336 | 0.393 | 0.210 | 0.242 |
| ProSST (K=2048) | 0.507 | 0.534 | 0.757 | 0.778 | 0.777 | 0.791 | 0.398 | 0.416 | 0.236 | 0.249 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ESM-IF1 | 0.421 | 0.474 | 0.747 | 0.777 | 0.729 | 0.759 | 0.328 | 0.372 | 0.223 | 0.239 |
| MIF-ST | 0.430 | 0.458 | 0.772 | 0.778 | 0.736 | 0.752 | 0.339 | 0.362 | 0.233 | 0.235 |
| ProteinMPNN (v_48_002) | 0.378 | 0.443 | 0.741 | 0.770 | 0.706 | 0.741 | 0.302 | 0.345 | 0.218 | 0.239 |
| ProteinMPNN (v_48_010) | 0.385 | 0.451 | 0.747 | 0.774 | 0.709 | 0.745 | 0.303 | 0.350 | 0.225 | 0.244 |
| ProteinMPNN (v_48_020) | 0.389 | 0.456 | 0.746 | 0.774 | 0.710 | 0.747 | 0.304 | 0.353 | 0.226 | 0.245 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| CARP-38M | 0.314 | 0.415 | 0.709 | 0.755 | 0.675 | 0.729 | 0.248 | 0.322 | 0.186 | 0.212 |
| CARP-600K | 0.159 | 0.354 | 0.649 | 0.740 | 0.590 | 0.694 | 0.127 | 0.271 | 0.144 | 0.194 |
| CARP-640M | 0.390 | 0.451 | 0.747 | 0.767 | 0.716 | 0.748 | 0.307 | 0.355 | 0.208 | 0.223 |
| CARP-76M | 0.360 | 0.436 | 0.728 | 0.759 | 0.699 | 0.740 | 0.282 | 0.339 | 0.196 | 0.214 |
| ESM-1b (mask) | 0.389 | 0.455 | 0.742 | 0.765 | 0.715 | 0.750 | 0.306 | 0.360 | 0.202 | 0.218 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ProGen2-B | 0.330 | 0.422 | 0.736 | 0.765 | 0.682 | 0.731 | 0.259 | 0.332 | 0.193 | 0.212 |
| ProGen2-L | 0.327 | 0.416 | 0.737 | 0.764 | 0.681 | 0.728 | 0.258 | 0.328 | 0.192 | 0.214 |
| ProGen2-M | 0.337 | 0.423 | 0.736 | 0.764 | 0.687 | 0.732 | 0.266 | 0.333 | 0.191 | 0.213 |
| ProGen2-S | 0.288 | 0.406 | 0.708 | 0.747 | 0.658 | 0.721 | 0.223 | 0.314 | 0.175 | 0.207 |
| ProGen2-XL | 0.351 | 0.419 | 0.755 | 0.772 | 0.694 | 0.730 | 0.275 | 0.329 | 0.198 | 0.207 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ESM3 | 0.185 | 0.226 | 0.837 | 0.842 | 0.598 | 0.619 | 0.147 | 0.187 | 0.219 | 0.219 |
| ProSST (K=1024) | 0.162 | 0.242 | 0.822 | 0.845 | 0.585 | 0.627 | 0.131 | 0.187 | 0.201 | 0.254 |
| ProSST (K=128) | 0.139 | 0.230 | 0.817 | 0.840 | 0.575 | 0.620 | 0.103 | 0.176 | 0.181 | 0.234 |
| ProSST (K=20) | 0.143 | 0.238 | 0.820 | 0.841 | 0.576 | 0.627 | 0.114 | 0.189 | 0.191 | 0.225 |
| ProSST (K=2048) | 0.213 | 0.249 | 0.840 | 0.847 | 0.614 | 0.630 | 0.170 | 0.199 | 0.228 | 0.246 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ESM-IF1 | 0.216 | 0.267 | 0.833 | 0.846 | 0.613 | 0.640 | 0.164 | 0.206 | 0.203 | 0.235 |
| MIF-ST | 0.196 | 0.230 | 0.837 | 0.842 | 0.599 | 0.618 | 0.141 | 0.172 | 0.219 | 0.229 |
| ProteinMPNN (v_48_002) | 0.191 | 0.244 | 0.830 | 0.844 | 0.598 | 0.626 | 0.149 | 0.183 | 0.199 | 0.230 |
| ProteinMPNN (v_48_010) | 0.218 | 0.256 | 0.836 | 0.848 | 0.611 | 0.631 | 0.163 | 0.199 | 0.229 | 0.253 |
| ProteinMPNN (v_48_020) | 0.221 | 0.271 | 0.835 | 0.851 | 0.611 | 0.638 | 0.164 | 0.194 | 0.225 | 0.266 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| CARP-38M | 0.073 | 0.158 | 0.816 | 0.831 | 0.543 | 0.587 | 0.057 | 0.120 | 0.195 | 0.206 |
| CARP-600K | 0.021 | 0.151 | 0.804 | 0.830 | 0.513 | 0.581 | 0.013 | 0.109 | 0.160 | 0.198 |
| CARP-640M | 0.157 | 0.206 | 0.830 | 0.838 | 0.584 | 0.609 | 0.128 | 0.152 | 0.192 | 0.212 |
| CARP-76M | 0.131 | 0.194 | 0.826 | 0.837 | 0.567 | 0.600 | 0.099 | 0.152 | 0.200 | 0.224 |
| ESM-1b (mask) | 0.166 | 0.211 | 0.831 | 0.838 | 0.589 | 0.612 | 0.137 | 0.162 | 0.208 | 0.215 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ProGen2-B | 0.124 | 0.194 | 0.823 | 0.837 | 0.564 | 0.598 | 0.101 | 0.159 | 0.184 | 0.210 |
| ProGen2-L | 0.120 | 0.188 | 0.823 | 0.835 | 0.560 | 0.598 | 0.091 | 0.147 | 0.187 | 0.210 |
| ProGen2-M | 0.115 | 0.182 | 0.825 | 0.835 | 0.558 | 0.595 | 0.078 | 0.133 | 0.189 | 0.206 |
| ProGen2-S | 0.091 | 0.192 | 0.815 | 0.838 | 0.546 | 0.600 | 0.069 | 0.157 | 0.174 | 0.223 |
| ProGen2-XL | 0.123 | 0.176 | 0.827 | 0.836 | 0.561 | 0.591 | 0.083 | 0.134 | 0.195 | 0.202 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ESM3 | 0.137 | 0.249 | 0.690 | 0.733 | 0.573 | 0.632 | 0.103 | 0.200 | 0.124 | 0.152 |
| ProSST (K=1024) | 0.204 | 0.275 | 0.673 | 0.727 | 0.608 | 0.645 | 0.157 | 0.219 | 0.125 | 0.153 |
| ProSST (K=128) | 0.179 | 0.264 | 0.671 | 0.725 | 0.596 | 0.640 | 0.139 | 0.212 | 0.127 | 0.152 |
| ProSST (K=20) | 0.161 | 0.259 | 0.669 | 0.721 | 0.586 | 0.637 | 0.122 | 0.212 | 0.126 | 0.155 |
| ProSST (K=2048) | 0.271 | 0.310 | 0.691 | 0.730 | 0.643 | 0.664 | 0.223 | 0.252 | 0.135 | 0.157 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ESM-IF1 | 0.285 | 0.318 | 0.720 | 0.755 | 0.648 | 0.666 | 0.226 | 0.258 | 0.160 | 0.171 |
| MIF-ST | 0.288 | 0.317 | 0.736 | 0.752 | 0.653 | 0.669 | 0.234 | 0.259 | 0.166 | 0.173 |
| ProteinMPNN (v_48_002) | 0.269 | 0.305 | 0.729 | 0.754 | 0.647 | 0.663 | 0.229 | 0.252 | 0.170 | 0.180 |
| ProteinMPNN (v_48_010) | 0.271 | 0.306 | 0.728 | 0.753 | 0.647 | 0.664 | 0.225 | 0.253 | 0.169 | 0.178 |
| ProteinMPNN (v_48_020) | 0.270 | 0.306 | 0.722 | 0.749 | 0.647 | 0.663 | 0.225 | 0.255 | 0.164 | 0.175 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| CARP-38M | 0.027 | 0.222 | 0.630 | 0.727 | 0.515 | 0.621 | 0.028 | 0.178 | 0.103 | 0.145 |
| CARP-600K | 0.006 | 0.210 | 0.614 | 0.722 | 0.504 | 0.615 | 0.013 | 0.170 | 0.099 | 0.143 |
| CARP-640M | 0.092 | 0.215 | 0.658 | 0.709 | 0.547 | 0.614 | 0.071 | 0.170 | 0.115 | 0.135 |
| CARP-76M | 0.050 | 0.230 | 0.637 | 0.728 | 0.527 | 0.625 | 0.040 | 0.181 | 0.109 | 0.151 |
| ESM-1b (mask) | 0.093 | 0.223 | 0.649 | 0.710 | 0.545 | 0.616 | 0.064 | 0.180 | 0.111 | 0.138 |
| Model | Spearman | NDCG | AUC | MCC | Top-recall | |||||
|---|---|---|---|---|---|---|---|---|---|---|
| Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | Raw | VRH | |
| ProGen2-B | 0.076 | 0.175 | 0.674 | 0.716 | 0.543 | 0.596 | 0.064 | 0.141 | 0.123 | 0.141 |
| ProGen2-L | 0.064 | 0.173 | 0.675 | 0.724 | 0.537 | 0.594 | 0.049 | 0.139 | 0.124 | 0.144 |
| ProGen2-M | 0.117 | 0.212 | 0.688 | 0.730 | 0.562 | 0.612 | 0.090 | 0.164 | 0.128 | 0.149 |
| ProGen2-S | 0.045 | 0.171 | 0.648 | 0.711 | 0.528 | 0.593 | 0.039 | 0.140 | 0.113 | 0.137 |
| ProGen2-XL | 0.162 | 0.235 | 0.715 | 0.744 | 0.588 | 0.627 | 0.128 | 0.190 | 0.146 | 0.153 |