Homo sapiens · seed P01116 · 189 aa · family defined as ≥30% identity to that seed · compiled 09 August 2026
Every figure here is counted over the whole family rather than quoted from one entry.
706 distinct constructs across 1,445 entries. 1,286 polymer entities differ from the UniProt canonical sequence in some way, 107 carry a recognised expression tag and 0 carry a fusion partner.
"Differs from canonical" is not the same as "engineered". The canonical sequence is the full gene product, so a secreted protein whose structures all start after its signal peptide counts every one of them as different: lysozyme's most-used construct, residues 19–147 on 1,239 entities, is simply the mature protein. Read the construct column below for what was actually done, rather than this count.
| Entities | Length | Best (Å) | Best entry | What was made |
|---|---|---|---|---|
| 62 | 166 | 1.26 | 1CTQ | residues 1-166 |
| 58 | 185 | 1.31 | 7G83 | residues 1-184 |
| 42 | 170 | 1.16 | 6TAN | residues 1-169; G12C, C51S, C80L +7 more |
| 35 | 167 | 1.20 | 8TBG | residues 1-166 |
| 26 | 167 | 1.65 | 6V9M | residues 1-166; Y64A |
| 25 | 170 | 1.16 | 8R7W | residues 1-169; G12D, R151G, E153D +4 more |
| 15 | 167 | 1.67 | 4HDO | residues 1-167 |
| 15 | 170 | 1.24 | 4OBE | residues 1-169; R151G, E153D, Q165K +3 more |
| 13 | 178 | 1.48 | 9HU8 | residues 1-177 |
| 12 | 170 | 0.95 | 9IAY | residues 1-169; C118S, R151G, E153D +4 more |
| 12 | 172 | 1.14 | 9G4B | residues 1-169; C118S |
| 11 | 169 | residues 1-169; G12V, C118S, R151G +5 more | ||
| 10 | 170 | 1.15 | 4LDJ | residues 1-169; G12C, R151G, E153D +4 more |
| 10 | 170 | 1.27 | 7RT1 | residues 1-169; G12D, C51S, C80L +7 more |
| 9 | 170 | 1.04 | 8AZX | residues 1-169; G12C, C118S, R151G +5 more |
| 9 | 183 | 1.50 | 6PGP | His7; residues 1-169; G12C, C51S, C80L +7 more |
| 9 | 185 | 2.17 | 4US0 | His6; TEV site; residues 1-166 |
| 9 | 186 | 1.18 | 6P0I | residues 1-187; 1 internal deletion; M180L, D182H, S183H +4 more |
| 9 | 189 | 2.61 | 9NI4 | 1 internal deletion; R151G, E153D, Q165K +15 more |
| 8 | 166 | 1.30 | 3OIW | residues 1-166; G12V |
| 8 | 169 | 1.49 | 6MS9 | residues 1-169; R151G, E153D, Q165K +3 more |
| 8 | 170 | 1.02 | 8AZZ | residues 1-169; G12V, C118S, R151G +5 more |
| 8 | 170 | 1.09 | 8AZY | residues 1-169; G12D, C118S, R151G +5 more |
| 8 | 177 | 1.34 | 9IFK | residues 1-177 |
| 8 | 193 | 1.75 | 6R3V | matches the canonical sequence |
Showing the 25 most-used of 706.
Columns where the wild-type residue still dominates but a real minority carries something else, which is a different question from "what varies across species".
| Oligomeric state | Chains | Entries | Share |
|---|---|---|---|
| monomeric | 1 | 710 | 49.1% |
| dimeric | 2 | 506 | 35.0% |
| trimeric | 3 | 110 | 7.6% |
| tetrameric | 4 | 75 | 5.2% |
| hexameric | 6 | 15 | 1.0% |
| pentameric | 5 | 9 | 0.6% |
| octameric | 8 | 8 | 0.6% |
| heptameric | 7 | 4 | 0.3% |
777 entries have the depositor's assembly corroborated by PISA, 627 carry the depositor's word alone and 41 were assigned by PISA where the depositor gave none. The middle figure is not a disagreement: PISA may have returned nothing or never run. 70 entries carry more than one assembly with different chain counts, so they have no single answer to quote: 1DOA, 1FOE, 1NF3, 1NVU, 1NVV, 1NVW, 1S8F, 1TU4, 1YU9, 1Z08, 1Z0A, 1ZBD, 2ATX, 2DFK, 2IEY, 2IEZ, 2IF0, 2IL1, 2NZJ, 2OCY.
| Domain | Source | Span (seed) | Chains |
|---|---|---|---|
| P-loop containing nucleotide triphosphate hydrolases | CATH 3.40.50.300 | 4–171 | 950 |
| Ras-like P-loop GTPases | SCOP2B 8019405 | 1–166 | 223 |
| Ras-like P-loop GTPases | SCOP2B 8057488 | 1–167 | 16 |
| Ras-like P-loop GTPases | SCOP2B 8060856 | 2–165 | 462 |
| Ras-like P-loop GTPases | SCOP2B 8057492 | 2–168 | 21 |
| Ras-like P-loop GTPases | SCOP2B 8057406 | 3–178 | 49 |
| Ras-like P-loop GTPases | SCOP2B 8057416 | 3–184 | 44 |
| Ras-like P-loop GTPases | SCOP2B 8054292 | 4–171 | 16 |
| Ras-like P-loop GTPases | SCOP2B 8057414 | 9–185 | 121 |
| Ras-like P-loop GTPases | SCOP2B 8089844 | 11–179 | 19 |
| Ras-like P-loop GTPases | SCOP2B 8057254 | 12–178 | 28 |
| Ras-like P-loop GTPases | SCOP2B 8057282 | 15–183 | 16 |
| Component | Class | Name | Entries | Best (Å) |
|---|---|---|---|---|
| MG | ion | Magnesium Ion | 1,125 | 0.95 |
| GDP | cofactor | Guanosine-5'-Diphosphate | 615 | 0.95 |
| GNP | cofactor | Phosphoaminophosphonic Acid-Guanylate Ester | 455 | 1.00 |
| GOL | cryoprotectant | Glycerol | 130 | 1.05 |
| CA | ion | Calcium Ion | 122 | 1.18 |
| GTP | cofactor | Guanosine-5'-Triphosphate | 97 | 1.19 |
| EDO | cryoprotectant | 1,2-Ethanediol | 96 | 0.95 |
| FMT | buffer | Formic Acid | 89 | 1.31 |
| SO4 | ion | Sulfate Ion | 86 | 1.12 |
| DMS | cryoprotectant | Dimethyl Sulfoxide | 65 | 1.31 |
| CL | ion | Chloride Ion | 57 | 1.08 |
| GSP | cofactor | 5'-Guanosine-Diphosphate-Monothiophosphate | 49 | 1.40 |
| NA | ion | Sodium Ion | 44 | 1.18 |
| GCP | ligand | Phosphomethylphosphonic Acid Guanylate Ester | 37 | 1.12 |
| ZN | ion | Zinc Ion | 37 | 1.81 |
| PO4 | ion | Phosphate Ion | 35 | 1.60 |
| ACT | cryoprotectant | Acetate Ion | 29 | 1.18 |
| AF3 | ligand | Aluminum Fluoride | 21 | 1.42 |
| PEG | cryoprotectant | Di(Hydroxyethyl)ether | 21 | 1.01 |
| 6IC | ligand | 4-(4-[(1r,5s)-3,8-Diazabicyclo[3.2.1]octan-3-Yl]-8-Fluoro-2-{[(2 | 19 | 1.23 |
Parsed from the free text 1,274 depositors typed into
_exptl_crystal_grow.pdbx_details, out of 1,290
entries that recorded anything at all.
Median pH 7.0
(range 1.0 to 10.5).
1,445 entries carry a wwPDB validation report: 613 clean, 574 worth a check and 258 with something to explain. Median clashscore 4.68, median RSRZ outliers 4.25%, median R-free minus R-work 0.037. 1,391 have released structure factors.
| Organism | Entries | Best (Å) | Ligand-bound | Seed covered |
|---|---|---|---|---|
| Homo sapiens | 1,313 | 0.95 | 1291 | 100% |
| Mus musculus | 35 | 1.20 | 34 | 96% |
| Saccharomyces cerevisiae | 20 | 1.35 | 12 | 100% |
| Rattus norvegicus | 15 | 1.00 | 15 | 96% |
| Arabidopsis thaliana | 12 | 1.53 | 10 | 94% |
| Saccharomyces cerevisiae S288C | 9 | 1.70 | 7 | 86% |
| Hordeum vulgare | 5 | 1.33 | 5 | 93% |
| Drosophila melanogaster | 5 | 1.39 | 3 | 87% |
| Leishmania donovani | 4 | 1.80 | 3 | 84% |
| Saguinus oedipus | 3 | 1.50 | 3 | 85% |
| Candidatus Thorarchaeota archaeon SMTZ1-45 | 3 | 1.50 | 3 | 84% |
| Avena sativa | 3 | 1.60 | 3 | 84% |
189 residues. Every identity figure in this document is measured against this sequence.
One record per paper, not per entry.
| Year | Citation |
|---|---|
| 2026 | Expanding Addressable KRAS Mutations through the Structure- and Property-Based Design of Dual-State (GDP/GTP), Reversible Pan-KRAS Inhibitors. J.Med.Chem. doi:10.1021/acs.jmedchem.6c01325 |
| 2026 | Selective Inhibition of KRASG13C Reveals an Increased Dependence on Wild-Type RAS Isoforms in Codon 13 RAS-Mutant Cancers. Cancer Discov doi:10.1158/2159-8290.CD-25-0886 |
| 2026 | From KRAS G12D to Pan-KRAS Inhibitors─A Journey Enabled by Synthetic Innovation and Structure-Based Drug Design. J.Med.Chem. doi:10.1021/acs.jmedchem.5c03662 |
| 2026 | Nucleotide-dependent switching and RIPb effector recognition of the barley susceptibility factor RACB. Commun Biol doi:10.1038/s42003-026-10316-7 |
| 2026 | Discovery of BBO-11818, a Potent and Selective Noncovalent Inhibitor of (ON) and (OFF) KRAS with Activity against Multiple Oncogenic Mutants. Cancer Discov doi:10.1158/2159-8290.CD-25-1280 |
| 2026 | Structure of SHOC2-KRAS-PP1C complex reveals RAS isoform-specific determinants and insights into targeting complex assembly by RAS inhibitors. Nat Commun doi:10.1038/s41467-026-68319-1 |
| 2026 | Targeting the H/KRAS alpha 4-beta 6-alpha 5 Allosteric Lobe with Macrocyclic Peptides. Acs Med.Chem.Lett. doi:10.1021/acsmedchemlett.6c00078 |
| 2026 | Discovering Targetable Conformation of RhoA Mutant by Integrating Native Mass Spectrometry, Ultraviolet Photodissociation, and X-ray Diffraction. J.Am.Chem.Soc. doi:10.1021/jacs.5c20067 |
| 2026 | Optimization of Covalent Warhead Trajectory for KRAS G12C Active-State Inhibition. J.Med.Chem. doi:10.1021/acs.jmedchem.5c03306 |
| 2026 | Optimization of Covalent 6-Cyanoquinazoline KRAS G12C Inhibitors for the Treatment of Solid Tumors. J.Med.Chem. doi:10.1021/acs.jmedchem.5c03610 |
| 2026 | Systematic cysteine scanning identifies a druggable pocket in oncogenic KRAS. Cell Chem Biol doi:10.1016/j.chembiol.2026.01.007 |
| 2026 | Covalent inhibitor design confers activity against both GDP- and GTP-bound forms of KRAS G12C. Nat Commun doi:10.1038/s41467-026-69003-0 |
| 2026 | Structure-Guided Development of NRAS G12D Inhibitors Based on a 5‐Azaindole Core. Acs Med.Chem.Lett. doi:10.1021/acsmedchemlett.5c00647 |
| 2026 | Disrupted molecular glue complex drives RAS inhibitor resistance. Cell doi:10.1016/j.cell.2026.03.031 |
| 2026 | Targeting Multiple KRAS Mutations with High-Affinity Macrocyclic Inhibitors: From Discovery to Preclinical Validation. J.Med.Chem. doi:10.1021/acs.jmedchem.6c00127 |
| 2026 | Structure-Activity Relationship Analysis of Macrocyclic Peptide RAS Inhibitors: Spotlight on the Solvent-Exposed Region. Acs Med.Chem.Lett. doi:10.1021/acsmedchemlett.6c00122 |
| 2026 | Analysis of structure and stability of Leishmania donovani Rab5a and Rab5b. Int.J.Biol.Macromol. doi:10.1016/j.ijbiomac.2026.152349 |
| 2026 | The cytokinesis regulator RacGAP1 is a Rac1-specific GAP on membranes. Protein Sci. doi:10.1002/pro.70488 |
| 2026 | Discovery and Characterization of Divarasib (GDC-6036), a Potent Covalent Inhibitor of KRAS G12C. J.Med.Chem. doi:10.1021/acs.jmedchem.5c02272 |
| 2026 | Discovery and Optimization of a Potent, Efficacious, and Brain-Penetrant Inhibitor of KRAS G12C. J.Med.Chem. doi:10.1021/acs.jmedchem.5c02279 |
| 2026 | An evolutionarily conserved salt bridge stabilizes the active site for GTP hydrolysis in Rho GTPases. J.Biol.Chem. doi:10.1016/j.jbc.2026.111260 |
| 2026 | Structures of the PI3K alpha /KRas complex on lipid bilayers reveal molecular mechanisms of PI3K alpha activation. Mol.Cell doi:10.1016/j.molcel.2026.06.010 |
| 2026 | Structural basis of fungal beta-1,3-glucan synthase inhibition by caspofungin. Nature doi:10.1038/s41586-026-10409-7 |
| 2026 | A novel RAB5 binding site in human VPS34-CII that is likely the primordial site in eukaryotic evolution. Elife doi:10.7554/eLife.110040 |
| 2026 | Protein Engineering-Enabled Cryo-EM Investigation of Small GTPases. J.Mol.Biol. doi:10.1016/j.jmb.2026.169860 |
| 2026 | Cyclized Peptide Inhibitors of the Small G Protein Cdc42 Mimic Binding of Effector Proteins. Biochemistry doi:10.1021/acs.biochem.5c00616 |
| 2025 | Discovery of BI-2493, a Pan-KRAS Inhibitor Showing In Vivo Efficacy. J.Med.Chem. doi:10.1021/acs.jmedchem.5c00576 |
| 2025 | Revealing Functional Hotspots: Temperature-Dependent Crystallography of K-RAS Highlights Allosteric and Druggable Sites. Biorxiv doi:10.1101/2025.02.27.639303 |
| 2025 | A neomorphic protein interface catalyzes covalent inhibition of RAS G12D aspartic acid in tumors. Science doi:10.1126/science.ads0239 |
| 2025 | Discovery of Daraxonrasib (RMC-6236), a Potent and Orally Bioavailable RAS(ON) Multi-selective, Noncovalent Tri-complex Inhibitor for the Treatment of Patients with Multiple RAS-Addicted Cancers. J.Med.Chem. doi:10.1021/acs.jmedchem.4c02314 |
| 2025 | Reversible Small Molecule Multivariant Ras Inhibitors Display Tunable Affinity for the Active and Inactive Forms of Ras. J.Med.Chem. doi:10.1021/acs.jmedchem.4c02929 |
| 2025 | Identification of Structurally Novel KRAS G12C Inhibitors through Covalent DNA-Encoded Library Screening. J.Med.Chem. doi:10.1021/acs.jmedchem.4c03071 |
| 2025 | Discovery of Elironrasib (RMC-6291), a Potent and Orally Bioavailable, RAS(ON) G12C-Selective, Covalent Tricomplex Inhibitor for the Treatment of Patients with RAS G12C-Addicted Cancers. J.Med.Chem. doi:10.1021/acs.jmedchem.4c02313 |
| 2025 | Hierarchical folding-upon-binding of an intrinsically disordered protein. Nat Commun doi:10.1038/s41467-025-66420-5 |
| 2025 | Design, Structure Optimization, and Preclinical Characterization of JAB-21822, a Covalent Inhibitor of KRAS G12C. J.Med.Chem. doi:10.1021/acs.jmedchem.4c02939 |
| 2025 | Promise and Challenge of beta-Lactone Electrophiles to Target Aspartate 12 of Mutant KRAS G12D . J.Med.Chem. doi:10.1021/acs.jmedchem.5c01214 |
| 2025 | Discovery of INCB159020, an Orally Bioavailable KRAS G12D Inhibitor. J.Med.Chem. doi:10.1021/acs.jmedchem.4c02662 |
| 2025 | Discovery of KRAS(G12D) selective degrader ASP3082. Commun Chem doi:10.1038/s42004-025-01662-4 |
| 2025 | Structural insights into isoform-specific RAS-PI3K alpha interactions and the role of RAS in PI3K alpha activation. Nat Commun doi:10.1038/s41467-024-55766-x |
| 2025 | Small-Molecule KRAS Inhibitors by Tyrosine Covalent Bond Formation. Chemmedchem doi:10.1002/cmdc.202400624 |