9ED4: Target of rapamycin complex subunit LST8
A composite map of mTORC1-Rag-Ragultor-4EBP1 on membrane. Determined by electron microscopy at 3.23 Å resolution. Released 10 Sept 2025.
- Method
- Electron microscopy
- Resolution
- 3.23 Å
- Organism
- Homo sapiens
- Chains
- 24
- Atoms
- 80,770
- Mol. weight
- 1312.89 kDa
- Ligands
- GSP, MG, ANP, IHP
- Released
- 10 Sept 2025
Explore 9ED4 in 3D
Show helices and sheets
RCSB PDB
PDBe
Secondary structure: helices and β-sheets
9ED4 contains 513 α-helices and 274 β-strands across 22 chains. Residue ranges use author residue numbering, as in the PDB file, from PDBe. To see them in 3D, choose the Cartoon representation with Secondary structure coloring: helices, sheets and coils get different colors.
Chain A: 149 helices, 14 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 16-29 | 14 | |
| α-helix | 34-50 | 17 | |
| α-helix | 57-76 | 20 | |
| α-helix | 80-93 | 14 | |
| α-helix | 101-114 | 14 | |
| α-helix | 120-135 | 16 | |
| α-helix | 141-156 | 16 | |
| α-helix | 162-178 | 17 | |
| α-helix | 180-184 | 5 | |
| α-helix | 187-198 | 12 | |
| α-helix | 203-223 | 21 | |
| α-helix | 233-245 | 13 | |
| α-helix | 248-250 | 3 | |
| α-helix | 257-275 | 19 | |
| α-helix | 277-303 | 27 | |
| α-helix | 360-368 | 9 | |
| α-helix | 370-380 | 11 | |
| α-helix | 386-402 | 17 | |
| α-helix | 404-410 | 7 | |
| α-helix | 413-423 | 11 | |
| α-helix | 430-443 | 14 | |
| α-helix | 452-462 | 11 | |
| α-helix | 464-465 | 2 | |
| α-helix | 480-493 | 14 | |
| α-helix | 495-497 | 3 | |
| α-helix | 498-511 | 14 | |
| α-helix | 515-527 | 13 | |
| α-helix | 529-531 | 3 | |
| α-helix | 532-547 | 16 | |
| α-helix | 579-591 | 13 | |
| α-helix | 599-605 | 7 | |
| α-helix | 606-610 | 5 | |
| α-helix | 616-629 | 14 | |
| α-helix | 631-638 | 8 | |
| α-helix | 641-643 | 3 | |
| α-helix | 646-665 | 20 | |
| α-helix | 669-678 | 10 | |
| α-helix | 681-683 | 3 | |
| α-helix | 684-687 | 4 | |
| α-helix | 690-699 | 10 | |
| α-helix | 705-719 | 15 | |
| α-helix | 723-743 | 21 | |
| α-helix | 747-763 | 17 | |
| α-helix | 765-768 | 4 | |
| α-helix | 772-783 | 12 | |
| α-helix | 793-808 | 16 | |
| α-helix | 812-814 | 3 | |
| α-helix | 817-829 | 13 | |
| α-helix | 837-851 | 15 | |
| α-helix | 856-860 | 5 | |
| α-helix | 863-874 | 12 | |
| α-helix | 878-891 | 14 | |
| β-strand | 894 | 1 | 9 |
| α-helix | 896-903 | 8 | |
| α-helix | 908-911 | 4 | |
| α-helix | 931-938 | 8 | |
| α-helix | 943-960 | 18 | |
| α-helix | 962-964 | 3 | |
| α-helix | 965-982 | 18 | |
| α-helix | 983-989 | 7 | |
| α-helix | 990-1003 | 14 | |
| α-helix | 1006-1023 | 18 | |
| α-helix | 1024-1027 | 4 | |
| α-helix | 1028-1041 | 14 | |
| α-helix | 1046-1062 | 17 | |
| α-helix | 1067-1070 | 4 | |
| α-helix | 1071-1074 | 4 | |
| α-helix | 1075-1083 | 9 | |
| α-helix | 1088-1090 | 3 | |
| α-helix | 1091-1101 | 11 | |
| α-helix | 1104-1107 | 4 | |
| α-helix | 1111-1123 | 13 | |
| α-helix | 1129-1143 | 15 | |
| α-helix | 1149-1165 | 17 | |
| α-helix | 1170-1184 | 15 | |
| α-helix | 1186-1202 | 17 | |
| α-helix | 1207-1211 | 5 | |
| α-helix | 1262-1268 | 7 | |
| α-helix | 1271-1273 | 3 | |
| α-helix | 1277-1294 | 18 | |
| α-helix | 1298-1302 | 5 | |
| α-helix | 1304-1309 | 6 | |
| α-helix | 1311-1316 | 6 | |
| α-helix | 1318-1326 | 9 | |
| α-helix | 1331-1347 | 17 | |
| α-helix | 1351-1366 | 16 | |
| α-helix | 1380-1390 | 11 | |
| α-helix | 1393-1406 | 14 | |
| α-helix | 1410-1422 | 13 | |
| α-helix | 1426-1439 | 14 | |
| α-helix | 1444-1446 | 3 | |
| α-helix | 1447-1453 | 7 | |
| α-helix | 1456-1467 | 12 | |
| α-helix | 1474-1487 | 14 | |
| α-helix | 1490-1500 | 11 | |
| α-helix | 1501-1503 | 3 | |
| α-helix | 1506-1523 | 18 | |
| α-helix | 1526-1535 | 10 | |
| α-helix | 1541-1553 | 13 | |
| α-helix | 1557-1579 | 23 | |
| α-helix | 1583-1607 | 25 | |
| α-helix | 1612-1625 | 14 | |
| β-strand | 1626 | 1 | 9 |
| α-helix | 1630-1640 | 11 | |
| α-helix | 1650-1661 | 12 | |
| α-helix | 1666-1677 | 12 | |
| α-helix | 1681-1683 | 3 | |
| α-helix | 1688-1690 | 3 | |
| α-helix | 1694-1706 | 13 | |
| α-helix | 1710-1730 | 21 | |
| α-helix | 1737-1762 | 26 | |
| α-helix | 1768-1782 | 15 | |
| α-helix | 1787-1812 | 26 | |
| α-helix | 1870-1895 | 26 | |
| α-helix | 1900-1913 | 14 | |
| α-helix | 1917-1929 | 13 | |
| α-helix | 1933-1938 | 6 | |
| α-helix | 1939-1944 | 6 | |
| α-helix | 1951-1967 | 17 | |
| α-helix | 1969-1980 | 12 | |
| α-helix | 1985-2020 | 36 | |
| α-helix | 2023-2036 | 14 | |
| α-helix | 2037-2041 | 5 | |
| α-helix | 2044-2060 | 17 | |
| α-helix | 2065-2070 | 6 | |
| α-helix | 2071-2075 | 5 | |
| α-helix | 2076-2091 | 16 | |
| α-helix | 2094-2114 | 21 | |
| α-helix | 2115-2117 | 3 | |
| β-strand | 2120-2122 | 3 | 10 |
| α-helix | 2123-2126 | 4 | |
| α-helix | 2128-2132 | 5 | |
| β-strand | 2139 | 1 | 11 |
| α-helix | 2140 | 1 | |
| α-helix | 2149-2151 | 3 | |
| β-strand | 2152 | 1 | 11 |
| β-strand | 2153-2156 | 4 | 10 |
| β-strand | 2159-2162 | 4 | 10 |
| β-strand | 2170-2176 | 7 | 10 |
| β-strand | 2181-2187 | 7 | 10 |
| α-helix | 2192-2211 | 20 | |
| α-helix | 2213-2218 | 6 | |
| α-helix | 2221-2223 | 3 | |
| β-strand | 2227-2229 | 3 | 10 |
| β-strand | 2235-2238 | 4 | 10 |
| β-strand | 2243-2245 | 3 | 12 |
| α-helix | 2246-2256 | 11 | |
| α-helix | 2263-2271 | 9 | |
| α-helix | 2275-2277 | 3 | |
| α-helix | 2280-2292 | 13 | |
| α-helix | 2298-2305 | 8 | |
| α-helix | 2310-2334 | 25 | |
| α-helix | 2341-2343 | 3 | |
| β-strand | 2344-2347 | 4 | 12 |
| β-strand | 2353-2355 | 3 | 12 |
| α-helix | 2381-2386 | 6 | |
| α-helix | 2395-2409 | 15 | |
| α-helix | 2411-2422 | 12 | |
| α-helix | 2426-2428 | 3 | |
| α-helix | 2491-2509 | 19 | |
| α-helix | 2521-2533 | 13 | |
| α-helix | 2535-2539 | 5 | |
| α-helix | 2543-2545 | 3 | |
Chain B: 3 helices, 28 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 14-19 | 6 | 1 |
| β-strand | 23-27 | 5 | 1 |
| β-strand | 37-39 | 3 | 1 |
| β-strand | 45-50 | 6 | 2 |
| β-strand | 57-68 | 12 | 2 |
| α-helix | 75-77 | 3 | |
| β-strand | 79-87 | 9 | 2 |
| β-strand | 88-93 | 6 | 3 |
| β-strand | 99-104 | 6 | 3 |
| β-strand | 108-113 | 6 | 3 |
| β-strand | 123-126 | 4 | 3 |
| β-strand | 131-136 | 6 | 4 |
| β-strand | 142-147 | 6 | 4 |
| β-strand | 152-156 | 5 | 4 |
| β-strand | 162-165 | 4 | 4 |
| β-strand | 173-178 | 6 | 5 |
| β-strand | 184-189 | 6 | 5 |
| β-strand | 194-198 | 5 | 5 |
| α-helix | 199-200 | 2 | |
| β-strand | 211-212 | 2 | 5 |
| β-strand | 216 | 1 | 5 |
| β-strand | 223-228 | 6 | 6 |
| β-strand | 234-239 | 6 | 6 |
| β-strand | 243-248 | 6 | 6 |
| β-strand | 257-259 | 3 | 6 |
| α-helix | 265-267 | 3 | |
| β-strand | 275-278 | 4 | 7 |
| β-strand | 284-289 | 6 | 7 |
| β-strand | 293-298 | 6 | 7 |
| β-strand | 304-308 | 5 | 7 |
| β-strand | 315-321 | 7 | 1 |
Chain C: 56 helices, 39 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 22-25 | 4 | |
| α-helix | 27-28 | 2 | |
| α-helix | 33-36 | 4 | |
| α-helix | 44-46 | 3 | |
| β-strand | 57-64 | 8 | 19 |
| β-strand | 85 | 1 | 20 |
| β-strand | 88 | 1 | 20 |
| α-helix | 90-92 | 3 | |
| α-helix | 95-111 | 17 | |
| β-strand | 118-123 | 6 | 19 |
| β-strand | 126 | 1 | 21 |
| α-helix | 127-141 | 15 | |
| β-strand | 145-151 | 7 | 19 |
| α-helix | 156-159 | 4 | |
| β-strand | 163-167 | 5 | 21 |
| β-strand | 174-178 | 5 | 21 |
| α-helix | 179-185 | 7 | |
| β-strand | 190-195 | 6 | 19 |
| α-helix | 199-222 | 24 | |
| α-helix | 227-230 | 4 | |
| α-helix | 233-236 | 4 | |
| β-strand | 241-244 | 4 | 19 |
| α-helix | 262-268 | 7 | |
| α-helix | 270-280 | 11 | |
| α-helix | 292-296 | 5 | |
| α-helix | 308-327 | 20 | |
| α-helix | 330-333 | 4 | |
| α-helix | 334-338 | 5 | |
| α-helix | 341-359 | 19 | |
| β-strand | 365-366 | 2 | 19 |
| α-helix | 370-372 | 3 | |
| α-helix | 377-394 | 18 | |
| α-helix | 395-399 | 5 | |
| α-helix | 403-406 | 4 | |
| α-helix | 408-420 | 13 | |
| α-helix | 428-430 | 3 | |
| α-helix | 433-439 | 7 | |
| α-helix | 446-457 | 12 | |
| α-helix | 461-470 | 10 | |
| α-helix | 473-479 | 7 | |
| α-helix | 487-499 | 13 | |
| α-helix | 505-512 | 8 | |
| α-helix | 515-522 | 8 | |
| α-helix | 531-543 | 13 | |
| α-helix | 547-555 | 9 | |
| α-helix | 558-565 | 8 | |
| α-helix | 571-585 | 15 | |
| α-helix | 589-597 | 9 | |
| α-helix | 600-605 | 6 | |
| α-helix | 606-609 | 4 | |
| α-helix | 613-625 | 13 | |
| α-helix | 635-649 | 15 | |
| α-helix | 650-653 | 4 | |
| α-helix | 657-673 | 17 | |
| α-helix | 675-691 | 17 | |
| α-helix | 781-788 | 8 | |
| α-helix | 798-799 | 2 | |
| α-helix | 807-819 | 13 | |
| α-helix | 824-842 | 19 | |
| α-helix | 919-922 | 4 | |
| α-helix | 924-926 | 3 | |
| α-helix | 959-964 | 6 | |
| α-helix | 965-968 | 4 | |
| α-helix | 984-1008 | 25 | |
| β-strand | 1015-1021 | 7 | 22 |
| β-strand | 1028-1031 | 4 | 23 |
| β-strand | 1038-1041 | 4 | 23 |
| β-strand | 1045-1049 | 5 | 23 |
| β-strand | 1056-1061 | 6 | 23 |
| β-strand | 1070-1076 | 7 | 24 |
| β-strand | 1083-1088 | 6 | 24 |
| β-strand | 1092-1097 | 6 | 24 |
| β-strand | 1107-1113 | 7 | 24 |
| β-strand | 1127-1132 | 6 | 25 |
| β-strand | 1137-1142 | 6 | 25 |
| β-strand | 1146-1151 | 6 | 25 |
| β-strand | 1156-1162 | 7 | 25 |
| β-strand | 1169-1175 | 7 | 26 |
| β-strand | 1180-1185 | 6 | 26 |
| β-strand | 1190-1194 | 5 | 26 |
| β-strand | 1203-1207 | 5 | 26 |
| β-strand | 1214-1219 | 6 | 27 |
| β-strand | 1227-1231 | 5 | 27 |
| β-strand | 1235-1239 | 5 | 27 |
| β-strand | 1246-1251 | 6 | 27 |
| β-strand | 1256-1261 | 6 | 28 |
| β-strand | 1268-1272 | 5 | 28 |
| β-strand | 1276-1280 | 5 | 28 |
| α-helix | 1285 | 1 | |
| β-strand | 1286-1287 | 2 | 28 |
| α-helix | 1288 | 1 | |
| α-helix | 1296-1298 | 3 | |
| β-strand | 1304-1309 | 6 | 22 |
| β-strand | 1316-1320 | 5 | 22 |
| β-strand | 1324-1329 | 6 | 22 |
Chain D: 12 helices, 11 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 7-14 | 8 | 13 |
| α-helix | 20-28 | 9 | |
| α-helix | 33-38 | 6 | |
| β-strand | 46-53 | 8 | 13 |
| β-strand | 56-62 | 7 | 13 |
| α-helix | 67-71 | 5 | |
| α-helix | 72-76 | 5 | |
| α-helix | 78-81 | 4 | |
| β-strand | 87-92 | 6 | 13 |
| α-helix | 99-116 | 18 | |
| β-strand | 121-126 | 6 | 13 |
| α-helix | 129-132 | 4 | |
| α-helix | 137-151 | 15 | |
| β-strand | 157-161 | 5 | 13 |
| α-helix | 168-178 | 11 | |
| α-helix | 184-197 | 14 | |
| β-strand | 202-206 | 5 | 14 |
| β-strand | 213-218 | 6 | 14 |
| α-helix | 227-244 | 18 | |
| β-strand | 249-256 | 8 | 14 |
| β-strand | 259-267 | 9 | 14 |
| β-strand | 270-276 | 7 | 14 |
| α-helix | 285-299 | 15 | |
Chain E: 11 helices, 10 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 63-68 | 6 | 29 |
| α-helix | 74-82 | 9 | |
| β-strand | 111-114 | 4 | 29 |
| α-helix | 133-136 | 4 | |
| β-strand | 139-145 | 7 | 29 |
| α-helix | 151-167 | 17 | |
| β-strand | 172-178 | 7 | 29 |
| α-helix | 187-206 | 20 | |
| β-strand | 213-218 | 6 | 29 |
| α-helix | 224-236 | 13 | |
| α-helix | 240-254 | 15 | |
| β-strand | 258-263 | 6 | 14 |
| β-strand | 269-272 | 4 | 14 |
| α-helix | 276-278 | 3 | |
| α-helix | 279-295 | 17 | |
| α-helix | 296-300 | 5 | |
| β-strand | 315-320 | 6 | 14 |
| β-strand | 325-331 | 7 | 14 |
| β-strand | 335-341 | 7 | 14 |
| α-helix | 343-346 | 4 | |
| α-helix | 349-366 | 18 | |
Chains F and W: 5 helices, 2 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 50-64 | 15 | |
| α-helix | 78-98 | 21 | |
| α-helix | 105-109 | 5 | |
| β-strand | 110 | 1 | 30 |
| α-helix | 115-117 | 3 | |
| α-helix | 126-145 | 20 | |
| β-strand | 155 | 1 | 15 |
Chains G and Q: 3 helices, 8 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 2 | 1 | 15 |
| α-helix | 4-15 | 12 | |
| β-strand | 21-25 | 5 | 16 |
| β-strand | 31-36 | 6 | 16 |
| α-helix | 42-61 | 20 | |
| β-strand | 69-73 | 5 | 16 |
| β-strand | 76 | 1 | 17 |
| β-strand | 79 | 1 | 17 |
| β-strand | 80-86 | 7 | 16 |
| β-strand | 89-95 | 7 | 16 |
| α-helix | 101-123 | 23 | |
Chain H: 5 helices, 6 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 4-12 | 9 | |
| α-helix | 13-15 | 3 | |
| β-strand | 21-25 | 5 | 16 |
| β-strand | 33-36 | 4 | 16 |
| α-helix | 46-50 | 5 | |
| α-helix | 52-60 | 9 | |
| β-strand | 68 | 1 | 17 |
| β-strand | 70-75 | 6 | 16 |
| β-strand | 79-84 | 6 | 16 |
| β-strand | 89-94 | 6 | 16 |
| α-helix | 101-121 | 21 | |
9 more chain groups are not listed. Open the entry in the viewer and use the sequence panel to see them.
Molecules and chains
| Molecule | Chains | Type | Length | Organism | UniProt |
|---|
| Target of rapamycin complex subunit LST8 | B, M | protein | 326 | Homo sapiens | Q9BVC4 (AlphaFold model) |
| GTP-binding protein Rheb | L, N | protein | 184 | Homo sapiens | Q15382 (AlphaFold model) |
| Serine/threonine-protein kinase mTOR | A, O | protein | 2549 | Homo sapiens | P42345 (AlphaFold model) |
| Ras-related GTP-binding protein A | D, P | protein | 313 | Homo sapiens | Q7L523 (AlphaFold model) |
| Ragulator complex protein LAMTOR2 | G, Q | protein | 125 | Homo sapiens | Q9Y2Q5 |
| Ragulator complex protein LAMTOR3 | H, R | protein | 124 | Homo sapiens | Q9UHA4 |
| Ragulator complex protein LAMTOR5 | J, S | protein | 91 | Homo sapiens | O43504 |
| Eukaryotic translation initiation factor 4E-binding protein 1 | K, T | protein | 118 | Homo sapiens | Q13541 |
| Regulatory-associated protein of mTOR | C, U | protein | 1335 | Homo sapiens | Q8N122 |
| Ras-related GTP-binding protein C | E, V | protein | 399 | Homo sapiens | Q9HB90 |
| Ragulator complex protein LAMTOR1 | F, W | protein | 161 | Homo sapiens | Q6IAA8 |
| Ragulator complex protein LAMTOR4 | I, X | protein | 99 | Homo sapiens | Q0VGL1 |
Sequence of entity 1 (B, M), FASTA
>9ED4_1 Target of rapamycin complex subunit LST8 (chains B, M)
MNTSPGTVGSDPVILATAGYDHTVRFWQAHSGICTRTVQHQDSQVNALEVTPDRSMIAAA
GYQHIRMYDLNSNNPNPIISYDGVNKNIASVGFHEDGRWMYTGGEDCTARIWDLRSRNLQ
CQRIFQVNAPINCVCLHPNQAELIVGDQSGAIHIWDLKTDHNEQLIPEPEVSITSAHIDP
DASYMAAVNSTGNCYVWNLTGGIGDEVTQLIPKTKIPAHTRYALQCRFSPDSTLLATCSA
DQTCKIWRTSNFSLMTELSIKSGNPGESSRGWMWGCAFSGDSQYIVTASSDNLARLWCVE
TGEIKREYGGHQKAVVCLAFNDSVLG
Sequence of entity 2 (L, N), FASTA
>9ED4_2 GTP-binding protein Rheb (chains L, N)
MPQSKSRKIAILGYRSVGKSSLTIQFVEGQFVDSYDPTIENTFTKLITVNGQEYHLQLVD
TAGQDEYSIFPQTYSIDINGYILVYSVTSIKSFEVIKVIHGKLLDMVGKVQIPIMLVGNK
KDLHMERVISYEEGKALAESWNAAFLESSAKENQTAVDVFRRIILEAEKMDGAASQGKSS
CSVM
Sequence of entity 3 (A, O), FASTA
>9ED4_3 Serine/threonine-protein kinase mTOR (chains A, O)
MLGTGPAAATTAATTSSNVSVLQQFASGLKSRNEETRAKAAKELQHYVTMELREMSQEES
TRFYDQLNHHIFELVSSSDANERKGGILAIASLIGVEGGNATRIGRFANYLRNLLPSNDP
VVMEMASKAIGRLAMAGDTFTAEYVEFEVKRALEWLGADRNEGRRHAAVLVLRELAISVP
TFFFQQVQPFFDNIFVAVWDPKQAIREGAVAALRACLILTTQREPKEMQKPQWYRHTFEE
AEKGFDETLAKEKGMNRDDRIHGALLILNELVRISSMEGERLREEMEEITQQQLVHDKYC
KDLMGFGTKPRHITPFTSFQAVQPQQSNALVGLLGYSSHQGLMGFGTSPSPAKSTLVESR
CCRDLMEEKFDQVCQWVLKCRNSKNSLIQMTILNLLPRLAAFRPSAFTDTQYLQDTMNHV
LSCVKKEKERTAAFQALGLLSVAVRSEFKVYLPRVLDIIRAALPPKDFAHKRQKAMQVDA
TVFTCISMLARAMGPGIQQDIKELLEPMLAVGLSPALTAVLYDLSRQIPQLKKDIQDGLL
KMLSLVLMHKPLRHPGMPKGLAHQLASPGLTTLPEASDVGSITLALRTLGSFEFEGHSLT
QFVRHCADHFLNSEHKEIRMEAARTCSRLLTPSIHLISGHAHVVSQTAVQVVADVLSKLL
VVGITDPDPDIRYCVLASLDERFDAHLAQAENLQALFVALNDQVFEIRELAICTVGRLSS
MNPAFVMPFLRKMLIQILTELEHSGIGRIKEQSARMLGHLVSNAPRLIRPYMEPILKALI
LKLKDPDPDPNPGVINNVLATIGELAQVSGLEMRKWVDELFIIIMDMLQDSSLLAKRQVA
LWTLGQLVASTGYVVEPYRKYPTLLEVLLNFLKTEQNQGTRREAIRVLGLLGALDPYKHK
VNIGMIDQSRDASAVSLSESKSSQDSSDYSTSEMLVNMGNLPLDEFYPAVSMVALMRIFR
DQSLSHHHTMVVQAITFIFKSLGLKCVQFLPQVMPTFLNVIRVCDGAIREFLFQQLGMLV
SFVKSHIRPYMDEIVTLMREFWVMNTSIQSTIILLIEQIVVALGGEFKLYLPQLIPHMLR
VFMHDNSPGRIVSIKLLAAIQLFGANLDDYLHLLLPPIVKLFDAPEAPLPSRKAALETVD
RLTESLDFTDYASRIIHPIVRTLDQSPELRSTAMDTLSSLVFQLGKKYQIFIPMVNKVLV
RHRINHQRYDVLICRIVKGYTLADEEEDPLIYQHRMLRSGQGDALASGPVETGPMKKLHV
STINLQKAWGAARRVSKDDWLEWLRRLSLELLKDSSSPSLRSCWALAQAYNPMARDLFNA
AFVSCWSELNEDQQDELIRSIELALTSQDIAEVTQTLLNLAEFMEHSDKGPLPLRDDNGI
VLLGERAAKCRAYAKALHYKELEFQKGPTPAILESLISINNKLQQPEAAAGVLEYAMKHF
GELEIQATWYEKLHEWEDALVAYDKKMDTNKDDPELMLGRMRCLEALGEWGQLHQQCCEK
WTLVNDETQAKMARMAAAAAWGLGQWDSMEEYTCMIPRDTHDGAFYRAVLALHQDLFSLA
QQCIDKARDLLDAELTAMAGESYSRAYGAMVSCHMLSELEEVIQYKLVPERREIIRQIWW
ERLQGCQRIVEDWQKILMVRSLVVSPHEDMRTWLKYASLCGKSGRLALAHKTLVLLLGVD
PSRQLDHPLPTVHPQVTYAYMKNMWKSARKIDAFQHMQHFVQTMQQQAQHAIATEDQQHK
QELHKLMARCFLKLGEWQLNLQGINESTIPKVLQYYSAATEHDRSWYKAWHAWAVMNFEA
VLHYKHQNQARDEKKKLRHASGANITNATTAATTAATATTTASTEGSNSESEAESTENSP
TPSPLQKKVTEDLSKTLLMYTVPAVQGFFRSISLSRGNNLQDTLRVLTLWFDYGHWPDVN
EALVEGVKAIQIDTWLQVIPQLIARIDTPRPLVGRLIHQLLTDIGRYHPQALIYPLTVAS
KSTTTARHNAANKILKNMCEHSNTLVQQAMMVSEELIRVAILWHEMWHEGLEEASRLYFG
ERNVKGMFEVLEPLHAMMERGPQTLKETSFNQAYGRDLMEAQEWCRKYMKSGNVKDLTQA
WDLYYHVFRRISKQLPQLTSLELQYVSPKLLMCRDLELAVPGTYDPNQPIIRIQSIAPSL
QVITSKQRPRKLTLMGSNGHEFVFLLKGHEDLRQDERVMQLFGLVNTLLANDPTSLRKNL
SIQRYAVIPLSTNSGLIGWVPHCDTLHALIRDYREKKKILLNIEHRIMLRMAPDYDHLTL
MQKVEVFEHAVNNTAGDDLAKLLWLKSPSSEVWFDRRTNYTRSLAVMSMVGYILGLGDRH
PSNLMLDRLSGKILHIDFGDCFEVAMTREKFPEKIPFRLTRMLTNAMEVTGLDGNYRITC
HTVMEVLREHKDSVMAVLEAFVYDPLLNWRLMDTNTKGNKRSRTRTDSYSAGQSVEILDG
VELGEPAHKKTGTTVPESIHSFIGDGLVKPEALNKKAIQIINRVRDKLTGRDFSHDDTLD
VPTQVELLIKQATSHENLCQCYIGWCPFW
Sequence of entity 4 (D, P), FASTA
>9ED4_4 Ras-related GTP-binding protein A (chains D, P)
MPNTAMKKKVLLMGKSGSGKTSMRSIIFANYIARDTRRLGATIDVEHSHVRFLGNLVLNL
WDCGGLDTFMENYFTSQRDNIFRNVEVLIYVFDVESRELEKDMHYYQSCLEAILQNSPDA
KIFCLVHKMDLVQEDQRDLIFKEREEDLRRLSRPLECACFRTSIWDETLYKAWSSIVYQL
IPNVQQLEMNLRNFAQIIEADEVLLFERATFLVISHYQCKEQRDVHRFEKISNIIKQFKL
SCSKLAASFQSMEVRNSNFAAFIDIFTSNTYVMVVMSDPSIPSAATLINIRNARKHFEKL
ERVDGPKHSLLMR
Sequence of entity 5 (G, Q), FASTA
>9ED4_5 Ragulator complex protein LAMTOR2 (chains G, Q)
MLRPKALTQVLSQANTGGVQSTLLLNNEGSLLAYSGYGDTDARVTAAIASNIWAAYDRNG
NQAFNEDNLKFILMDCMEGRVAITRVANLLLCMYAKETVGFGMLKAKAQALVQYLEEPLT
QVAAS
Sequence of entity 6 (H, R), FASTA
>9ED4_6 Ragulator complex protein LAMTOR3 (chains H, R)
MADDLKRFLYKKLPSVEGLHAIVVSDRDGVPVIKVANDNAPEHALRPGFLSTFALATDQG
SKLGLSKNKSIICYYNTYQVVQFNRLPLVVSFIASSSANTGLIVSLEKELAPLFEELRQV
VEVS
Sequence of entity 7 (J, S), FASTA
>9ED4_7 Ragulator complex protein LAMTOR5 (chains J, S)
MEATLEQHLEDTMKNPSIVGVLCTDSQGLNLGCRGTLSDEHAGVISVLAQQAAKLTSDPT
DIPVVCLESDNGNIMIQKHDGITVAVHKMAS
Sequence of entity 8 (K, T), FASTA
>9ED4_8 Eukaryotic translation initiation factor 4E-binding protein 1 (chains K, T)
MSGGSSCSQTPSRAIPATRRVVLGDGVQLPPGDYSTTPGGTLFSTTPGGTRIIYDRKFLM
ECRNSPVTKTPPRDLPTIPGVTSPSSDEPPMEASQSHLRNSPEDKRAGGEESQFEMDI
Sequence of entity 9 (C, U), FASTA
>9ED4_9 Regulatory-associated protein of mTOR (chains C, U)
MESEMLQSPLLGLGEEDEADLTDWNLPLAFMKKRHCEKIEGSKSLAQSWRMKDRMKTVSV
ALVLCLNVGVDPPDVVKTTPCARLECWIDPLSMGPQKALETIGANLQKQYENWQPRARYK
QSLDPTVDEVKKLCTSLRRNAKEERVLFHYNGHGVPRPTVNGEVWVFNKNYTQYIPLSIY
DLQTWMGSPSIFVYDCSNAGLIVKSFKQFALQREQELEVAAINPNHPLAQMPLPPSMKNC
IQLAACEATELLPMIPDLPADLFTSCLTTPIKIALRWFCMQKCVSLVPGVTLDLIEKIPG
RLNDRRTPLGELNWIFTAITDTIAWNVLPRDLFQKLFRQDLLVASLFRNFLLAERIMRSY
NCTPVSSPRLPPTYMHAMWQAWDLAVDICLSQLPTIIEEGTAFRHSPFFAEQLTAFQVWL
TMGVENRNPPEQLPIVLQVLLSQVHRLRALDLLGRFLDLGPWAVSLALSVGIFPYVLKLL
QSSARELRPLLVFIWAKILAVDSSCQADLVKDNGHKYFLSVLADPYMPAEHRTMTAFILA
VIVNSYHTGQEACLQGNLIAICLEQLNDPHPLLRQWVAICLGRIWQNFDSARWCGVRDSA
HEKLYSLLSDPIPEVRCAAVFALGTFVGNSAERTDHSTTIDHNVAMMLAQLVSDGSPMVR
KELVVALSHLVVQYESNFCTVALQFIEEEKNYALPSPATTEGGSLTPVRDSPCTPRLRSV
SSYGNIRAVATARSLNKSLQNLSLTEESGGAVAFSPGNLSTSSSASSTLGSPENEEHILS
FETIDKMRRASSYSSLNSLIGVSFNSVYTQIWRVLLHLAADPYPEVSDVAMKVLNSIAYK
ATVNARPQRVLDTSSLTQSAPASPTNKGVHIHQAGGSPPASSTSSSSLTNDVAKQPVSRD
LPSGRPGTTGPAGAQYTPHSHQFPRTRKMFDKGPEQTADDADDAAGHKSFISATVQTGFC
DWSARYFAQPVMKIPEEHDLESQIRKEREWRFLRNSRVRRQAQQVIQKGITRLDDQIFLN
RNPGVPSVVKFHPFTPCIAVADKDSICFWDWEKGEKLDYFHNGNPRYTRVTAMEYLNGQD
CSLLLTATDDGAIRVWKNFADLEKNPEMVTAWQGLSDMLPTTRGAGMVVDWEQETGLLMS
SGDVRIVRIWDTDREMKVQDIPTGADSCVTSLSCDSHRSLIVAGLGDGSIRVYDRRMALS
ECRVMTYREHTAWVVKASLQKRPDGHIVSVSVNGDVRIFDPRMPESVNVLQIVKGLTALD
IHPQADLIACGSVNQFTAIYNSSGELINNIKYYDGFMGQRVGAISCLAFHPHWPHLAVGS
NDYYISVYSVEKRVR
Sequence of entity 10 (E, V), FASTA
>9ED4_10 Ras-related GTP-binding protein C (chains E, V)
MSLQYGAEETPLAGSYGAADSFPKDFGYGVEEEEEEAAAAGGGVGAGAGGGCGPGGADSS
KPRILLMGLRRSGKNSIQKVVFHKMSPNETLFLESTNKIYKDDISNSSFVNFQIWDFPGQ
MDFFDPTFDYEMIFRGTGALIYVIDAQDDYMEALTRLHITVSKAYKVNPDMNFEVFIHKV
DGLSDDHKIETQRDIHQRANDDLADAGLEKLHLSFYLTSIYDHSIFEAFSKVVQKLIPQL
PTLENLLNIFISNSGIEKAFLFDVVSKIYIATDSSPVDMQSYELCCDMIDVVIDVSCIYG
LKEDGSGSAYDKESMAIIKLNNTTVLYLKEVTKFLALVCILREESFERKGLIDYNFHCFR
KAIHEVFEVGVTSHRSCGHQTSASSLKALTHNGTPRNAI
Sequence of entity 11 (F, W), FASTA
>9ED4_11 Ragulator complex protein LAMTOR1 (chains F, W)
MGCCYSSENEDSDQDREERKLLLDPSSPPTKALNGAEPNYHSLPSARTDEQALLSSILAK
TASNIIDVSAADSQGMEQHEYMDRARQYSTRLAVLSSSLTHWKKLPPLPSLTSQPHQVLA
SEPIPFSDLQQVSRIAAYAYSALSQIRVDAKEELVVQFGIP
Sequence of entity 12 (I, X), FASTA
>9ED4_12 Ragulator complex protein LAMTOR4 (chains I, X)
MTSALTQGLERIPDQLGYLVLSEGAVLASSGDLENDEQAASAISELVSTACGFRLHRGMN
VPFKRLSVVFGEHTLLVTVSGQRVFVVKRQNRGREPIDV
Ligands and cofactors
| ID | Name | Formula | Copies |
|---|
| GSP | 5'-guanosine-diphosphate-monothiophosphate | C10 H16 N5 O13 P3 S | 2 |
| MG | Magnesium ion | Mg | 8 |
| ANP | Phosphoaminophosphonic acid-adenylate ester | C10 H17 N6 O12 P3 | 2 |
| IHP | Inositol hexakisphosphate | C6 H18 O24 P6 | 2 |
| GTP | Guanosine-5'-triphosphate | C10 H16 N5 O14 P3 | 2 |
| GDP | Guanosine-5'-diphosphate | C10 H15 N5 O11 P2 | 2 |
Primary citation
Structural basis for mTORC1 activation on the lysosomal membrane. Cui, Z., Esposito, A., Napolitano, G. et al. Nature (2025) 647:536-543. DOI 10.1038/s41586-025-09545-3 · PubMed
Other PDB entries of the same protein (UniProt Q9BVC4 (AlphaFold model), which also has an AlphaFold model), best resolution first:
- 9T94 2.6 Å, cryo-EM structure of AKT phosphorylated mTOR complex 2, focused on a single protomer
- 9ZBK 2.6 Å, mTORC2 in complex with Akt1
- 8ERA 2.86 Å, RMC-5552 in complex with mTORC1 and FKBP12
- 9T93 2.86 Å, cryo-EM structure of autophosphorylated mTOR complex 2, focused on a single protomer
- 6ZWO 3.0 Å, cryo-EM structure of human mTOR complex 2, focused on one half
- 9T7J 3.0 Å, cryo-EM structure of AKT phosphorylated mTOR complex 2, overall refinement
- 9TDT 3.0 Å, cryo-EM structure of dephosphorylated mTOR complex 2, focused on a single protomer
- 5WBY 3.1 Å, Crystal structure of mTOR(deltaN)-mLST8-PRAS40(beta-strand) complex
- 9T92 3.1 Å, cryo-EM structure of autophosphorylated mTOR complex 2, overall refinement
- 9ED7 3.16 Å, Active state of mTOR on membrane
- 4JSN 3.2 Å, structure of mTORdeltaN-mLST8 complex
- 6ZWM 3.2 Å, cryo-EM structure of human mTOR complex 2, overall refinement
Browse structure collections
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