9E2X: DNA replication complex GINS protein PSF1
Cryo-EM structure of yeast CMG helicase stalled at G4-containing DNA template, state 2. Determined by electron microscopy at 3.5 Å resolution. Released 19 Mar 2025.
- Method
- Electron microscopy
- Resolution
- 3.5 Å
- Organisms
- Saccharomyces cerevisiae W303, synthetic construct
- Chains
- 15
- Atoms
- 49,316
- Mol. weight
- 965.18 kDa
- Ligands
- ATP, MG, ZN, ADP
- Released
- 19 Mar 2025
Explore 9E2X in 3D
Show helices and sheets
RCSB PDB
PDBe
Secondary structure: helices and β-sheets
9E2X contains 296 α-helices and 235 β-strands across 13 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 2: 28 helices, 30 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 176-178 | 3 | |
| α-helix | 183-187 | 5 | |
| α-helix | 194-199 | 6 | |
| α-helix | 201-217 | 17 | |
| β-strand | 219 | 1 | 14 |
| β-strand | 225 | 1 | 14 |
| α-helix | 226-236 | 11 | |
| β-strand | 241-244 | 4 | 15 |
| α-helix | 246-252 | 7 | |
| α-helix | 254-262 | 9 | |
| α-helix | 264-282 | 19 | |
| α-helix | 286-289 | 4 | |
| β-strand | 294-297 | 4 | 15 |
| β-strand | 303-304 | 2 | 16 |
| α-helix | 311-313 | 3 | |
| β-strand | 317-327 | 11 | 16 |
| β-strand | 331-334 | 4 | 16 |
| β-strand | 336-341 | 6 | 17 |
| β-strand | 347-352 | 6 | 17 |
| β-strand | 373-375 | 3 | 17 |
| β-strand | 382-391 | 10 | 16 |
| β-strand | 405-411 | 7 | 16 |
| β-strand | 423-434 | 12 | 16 |
| α-helix | 436-442 | 7 | |
| β-strand | 446 | 1 | 18 |
| β-strand | 447-458 | 12 | 16 |
| α-helix | 478-488 | 11 | |
| α-helix | 493-499 | 7 | |
| α-helix | 509-520 | 12 | |
| β-strand | 525-527 | 3 | 19 |
| β-strand | 531-533 | 3 | 19 |
| β-strand | 539-542 | 4 | 20 |
| α-helix | 550-559 | 10 | |
| β-strand | 563-567 | 5 | 20 |
| β-strand | 571 | 1 | 21 |
| α-helix | 573-577 | 5 | |
| β-strand | 580-582 | 3 | 22 |
| β-strand | 589-591 | 3 | 22 |
| α-helix | 595-598 | 4 | |
| β-strand | 603-607 | 5 | 20 |
| α-helix | 609-611 | 3 | |
| α-helix | 614-625 | 12 | |
| β-strand | 628-631 | 4 | 23 |
| β-strand | 638-641 | 4 | 23 |
| β-strand | 645-650 | 6 | 20 |
| α-helix | 663-666 | 4 | |
| α-helix | 671-676 | 6 | |
| β-strand | 679-682 | 4 | 20 |
| β-strand | 686 | 1 | 24 |
| α-helix | 689-705 | 17 | |
| α-helix | 740-755 | 16 | |
| α-helix | 760-773 | 14 | |
| β-strand | 777 | 1 | 25 |
| α-helix | 783-797 | 15 | |
| α-helix | 807-823 | 17 | |
| β-strand | 828 | 1 | 25 |
| α-helix | 831-848 | 18 | |
| α-helix | 851-860 | 10 | |
Chain 3: 24 helices, 28 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 20-34 | 15 | |
| α-helix | 36-51 | 16 | |
| β-strand | 95-99 | 5 | 26 |
| α-helix | 100-106 | 7 | |
| α-helix | 108-116 | 9 | |
| α-helix | 118-136 | 19 | |
| α-helix | 144-148 | 5 | |
| β-strand | 154-158 | 5 | 26 |
| β-strand | 165-166 | 2 | 27 |
| α-helix | 173-175 | 3 | |
| β-strand | 179-189 | 11 | 27 |
| α-helix | 190-192 | 3 | |
| β-strand | 193-203 | 11 | 27 |
| β-strand | 208-213 | 6 | 27 |
| β-strand | 234 | 1 | 28 |
| β-strand | 240 | 1 | 28 |
| β-strand | 242-259 | 18 | 27 |
| α-helix | 266 | 1 | |
| β-strand | 273-279 | 7 | 27 |
| β-strand | 291-301 | 11 | 27 |
| β-strand | 318-328 | 11 | 27 |
| α-helix | 344-355 | 12 | |
| α-helix | 359-366 | 8 | |
| α-helix | 375-386 | 12 | |
| β-strand | 391-392 | 2 | 29 |
| β-strand | 398-399 | 2 | 29 |
| β-strand | 405-409 | 5 | 30 |
| α-helix | 415-425 | 11 | |
| β-strand | 433 | 1 | 31 |
| β-strand | 445-447 | 3 | 32 |
| β-strand | 456-458 | 3 | 32 |
| α-helix | 463-465 | 3 | |
| β-strand | 468-471 | 4 | 30 |
| β-strand | 473 | 1 | 31 |
| α-helix | 475-477 | 3 | |
| α-helix | 480-492 | 13 | |
| β-strand | 494-499 | 6 | 33 |
| β-strand | 502-507 | 6 | 33 |
| β-strand | 510-516 | 7 | 30 |
| α-helix | 529-532 | 4 | |
| α-helix | 537-542 | 6 | |
| β-strand | 545-549 | 5 | 30 |
| α-helix | 555-570 | 16 | |
| β-strand | 622 | 1 | 34 |
| α-helix | 632-638 | 7 | |
| β-strand | 651 | 1 | 34 |
| α-helix | 653-666 | 14 | |
| β-strand | 670 | 1 | 35 |
| α-helix | 673-688 | 16 | |
| α-helix | 700-715 | 16 | |
| β-strand | 720 | 1 | 35 |
| α-helix | 723-737 | 15 | |
Chain 4: 29 helices, 35 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 177-180 | 4 | 36 |
| β-strand | 186 | 1 | 36 |
| α-helix | 187-199 | 13 | |
| β-strand | 202 | 1 | 37 |
| α-helix | 204-209 | 6 | |
| α-helix | 213-215 | 3 | |
| β-strand | 224 | 1 | 37 |
| α-helix | 225-236 | 12 | |
| β-strand | 240-244 | 5 | 38 |
| α-helix | 245-250 | 6 | |
| α-helix | 255-263 | 9 | |
| α-helix | 265-287 | 23 | |
| α-helix | 293-297 | 5 | |
| β-strand | 302-306 | 5 | 38 |
| α-helix | 319-321 | 3 | |
| β-strand | 325-335 | 11 | 39 |
| α-helix | 336-338 | 3 | |
| β-strand | 339-349 | 11 | 39 |
| β-strand | 355-359 | 5 | 39 |
| β-strand | 360-361 | 2 | 40 |
| β-strand | 364-365 | 2 | 40 |
| β-strand | 370 | 1 | 41 |
| β-strand | 379 | 1 | 41 |
| β-strand | 382-384 | 3 | 39 |
| α-helix | 386-388 | 3 | |
| β-strand | 390-400 | 11 | 39 |
| α-helix | 403-405 | 3 | |
| α-helix | 411-413 | 3 | |
| β-strand | 414-420 | 7 | 39 |
| α-helix | 421-423 | 3 | |
| β-strand | 432-442 | 11 | 39 |
| β-strand | 445 | 1 | 42 |
| β-strand | 453-454 | 2 | 42 |
| β-strand | 456 | 1 | 43 |
| α-helix | 457 | 1 | |
| β-strand | 458-468 | 11 | 39 |
| α-helix | 503-514 | 12 | |
| α-helix | 518-526 | 9 | |
| α-helix | 534-545 | 12 | |
| β-strand | 550 | 1 | 44 |
| β-strand | 558 | 1 | 44 |
| β-strand | 564-568 | 5 | 45 |
| α-helix | 575-584 | 10 | |
| β-strand | 590-591 | 2 | 45 |
| β-strand | 596 | 1 | 46 |
| α-helix | 598-602 | 5 | |
| β-strand | 603-608 | 6 | 47 |
| β-strand | 613-618 | 6 | 47 |
| α-helix | 620-623 | 4 | |
| β-strand | 628-631 | 4 | 45 |
| α-helix | 639-651 | 13 | |
| β-strand | 653-658 | 6 | 48 |
| β-strand | 661-666 | 6 | 48 |
| β-strand | 670-675 | 6 | 45 |
| α-helix | 688-691 | 4 | |
| α-helix | 696-701 | 6 | |
| β-strand | 704-708 | 5 | 45 |
| α-helix | 716-725 | 10 | |
| α-helix | 744-756 | 13 | |
| β-strand | 761-762 | 2 | 49 |
| α-helix | 764-779 | 16 | |
| α-helix | 795-811 | 17 | |
| β-strand | 816-817 | 2 | 49 |
| α-helix | 819-833 | 15 | |
Chain 5: 31 helices, 22 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 23-37 | 15 | |
| α-helix | 45-55 | 11 | |
| β-strand | 60-64 | 5 | 50 |
| α-helix | 65-71 | 7 | |
| α-helix | 73-81 | 9 | |
| α-helix | 83-102 | 20 | |
| α-helix | 133-135 | 3 | |
| β-strand | 136-140 | 5 | 50 |
| α-helix | 143-147 | 5 | |
| α-helix | 148-150 | 3 | |
| α-helix | 153-155 | 3 | |
| β-strand | 159-169 | 11 | 51 |
| β-strand | 173-183 | 11 | 51 |
| β-strand | 189-193 | 5 | 51 |
| β-strand | 241-259 | 19 | 51 |
| α-helix | 266 | 1 | |
| β-strand | 273-279 | 7 | 51 |
| α-helix | 280-282 | 3 | |
| β-strand | 291-301 | 11 | 51 |
| β-strand | 323 | 1 | 16 |
| β-strand | 325-335 | 11 | 51 |
| α-helix | 352-361 | 10 | |
| α-helix | 366-373 | 8 | |
| α-helix | 382-393 | 12 | |
| α-helix | 405-407 | 3 | |
| β-strand | 412-415 | 4 | 52 |
| α-helix | 422-432 | 11 | |
| β-strand | 436-440 | 5 | 52 |
| β-strand | 451-456 | 6 | 53 |
| β-strand | 461-466 | 6 | 53 |
| α-helix | 468-471 | 4 | |
| β-strand | 476-480 | 5 | 52 |
| α-helix | 487-499 | 13 | |
| β-strand | 518-523 | 6 | 52 |
| α-helix | 544-549 | 6 | |
| β-strand | 552-554 | 3 | 52 |
| α-helix | 562-577 | 16 | |
| α-helix | 596-609 | 14 | |
| β-strand | 613-614 | 2 | 54 |
| α-helix | 616-636 | 21 | |
| α-helix | 650-656 | 7 | |
| α-helix | 660-666 | 7 | |
| β-strand | 671-672 | 2 | 54 |
| α-helix | 674-684 | 11 | |
| α-helix | 685-689 | 5 | |
| α-helix | 690-693 | 4 | |
| α-helix | 707-721 | 15 | |
| β-strand | 727-729 | 3 | 55 |
| α-helix | 730-734 | 5 | |
| α-helix | 735-739 | 5 | |
| α-helix | 746-757 | 12 | |
| β-strand | 761-764 | 4 | 55 |
| β-strand | 769-772 | 4 | 55 |
Chain 6: 37 helices, 36 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 94-97 | 4 | 56 |
| α-helix | 99-101 | 3 | |
| α-helix | 103-104 | 2 | |
| α-helix | 106-121 | 16 | |
| β-strand | 124-125 | 2 | 57 |
| β-strand | 132-133 | 2 | 57 |
| α-helix | 135-146 | 12 | |
| β-strand | 150-154 | 5 | 58 |
| α-helix | 155-160 | 6 | |
| α-helix | 161-164 | 4 | |
| α-helix | 165-172 | 8 | |
| α-helix | 174-192 | 19 | |
| α-helix | 194-197 | 4 | |
| β-strand | 198 | 1 | 59 |
| α-helix | 259 | 1 | |
| β-strand | 260 | 1 | 59 |
| α-helix | 261 | 1 | |
| β-strand | 264-268 | 5 | 58 |
| β-strand | 273-274 | 2 | 18 |
| α-helix | 276-278 | 3 | |
| α-helix | 281-283 | 3 | |
| β-strand | 287-297 | 11 | 18 |
| α-helix | 298-300 | 3 | |
| β-strand | 301-311 | 11 | 18 |
| β-strand | 318-322 | 5 | 18 |
| α-helix | 323 | 1 | |
| β-strand | 344-346 | 3 | 18 |
| β-strand | 352-362 | 11 | 18 |
| α-helix | 369 | 1 | |
| α-helix | 373-375 | 3 | |
| β-strand | 376-382 | 7 | 18 |
| α-helix | 383-385 | 3 | |
| β-strand | 394-405 | 12 | 18 |
| α-helix | 407-410 | 4 | |
| β-strand | 437 | 1 | 60 |
| β-strand | 438 | 1 | 40 |
| β-strand | 448 | 1 | 60 |
| β-strand | 450-461 | 12 | 18 |
| α-helix | 501-507 | 7 | |
| α-helix | 510-521 | 12 | |
| α-helix | 525-532 | 8 | |
| α-helix | 541-552 | 12 | |
| β-strand | 557-558 | 2 | 61 |
| α-helix | 563 | 1 | |
| β-strand | 564-565 | 2 | 61 |
| α-helix | 566 | 1 | |
| β-strand | 571-575 | 5 | 62 |
| α-helix | 582-589 | 8 | |
| β-strand | 595-598 | 4 | 62 |
| β-strand | 604 | 1 | 63 |
| β-strand | 607 | 1 | 63 |
| β-strand | 611-614 | 4 | 64 |
| α-helix | 615 | 1 | |
| β-strand | 621-624 | 4 | 64 |
| α-helix | 627-630 | 4 | |
| β-strand | 635-638 | 4 | 62 |
| α-helix | 646-649 | 4 | |
| α-helix | 654-657 | 4 | |
| β-strand | 660-663 | 4 | 65 |
| β-strand | 664 | 1 | 21 |
| β-strand | 670-673 | 4 | 65 |
| β-strand | 677-682 | 6 | 62 |
| α-helix | 695-698 | 4 | |
| α-helix | 703-708 | 6 | |
| β-strand | 711-715 | 5 | 62 |
| α-helix | 721-737 | 17 | |
| α-helix | 748-758 | 11 | |
| β-strand | 765 | 1 | 66 |
| α-helix | 767-784 | 18 | |
| β-strand | 791 | 1 | 24 |
| α-helix | 797-813 | 17 | |
| β-strand | 819 | 1 | 66 |
| α-helix | 821-834 | 14 | |
Chain 7: 30 helices, 34 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 7-8 | 2 | 27 |
| α-helix | 14-27 | 14 | |
| α-helix | 62-71 | 10 | |
| β-strand | 78-81 | 4 | 67 |
| α-helix | 82-95 | 14 | |
| α-helix | 102-108 | 7 | |
| α-helix | 113-123 | 11 | |
| α-helix | 125-128 | 4 | |
| α-helix | 131-132 | 2 | |
| α-helix | 138-155 | 18 | |
| α-helix | 194-197 | 4 | |
| β-strand | 202-205 | 4 | 67 |
| α-helix | 209-214 | 6 | |
| β-strand | 224-225 | 2 | 43 |
| α-helix | 232-234 | 3 | |
| β-strand | 238-247 | 10 | 43 |
| β-strand | 252-262 | 11 | 43 |
| β-strand | 268-272 | 5 | 43 |
| β-strand | 277-278 | 2 | 42 |
| β-strand | 283 | 1 | 68 |
| α-helix | 287-291 | 5 | |
| α-helix | 296 | 1 | |
| β-strand | 297 | 1 | 68 |
| β-strand | 298-300 | 3 | 43 |
| β-strand | 306-316 | 11 | 43 |
| α-helix | 323 | 1 | |
| β-strand | 330-336 | 7 | 43 |
| β-strand | 347-357 | 11 | 43 |
| α-helix | 363-368 | 6 | |
| β-strand | 371 | 1 | 27 |
| β-strand | 374-383 | 10 | 43 |
| α-helix | 410-413 | 4 | |
| α-helix | 426-437 | 12 | |
| β-strand | 444 | 1 | 69 |
| β-strand | 449 | 1 | 69 |
| β-strand | 456-460 | 5 | 70 |
| β-strand | 462 | 1 | 71 |
| β-strand | 464 | 1 | 71 |
| α-helix | 467-476 | 10 | |
| β-strand | 480-484 | 5 | 70 |
| β-strand | 488 | 1 | 72 |
| β-strand | 491 | 1 | 72 |
| β-strand | 495-499 | 5 | 73 |
| β-strand | 506-510 | 5 | 73 |
| α-helix | 512-515 | 4 | |
| β-strand | 520-524 | 5 | 70 |
| α-helix | 526-528 | 3 | |
| α-helix | 531-542 | 12 | |
| β-strand | 546-547 | 2 | 74 |
| β-strand | 549 | 1 | 46 |
| β-strand | 556-557 | 2 | 74 |
| β-strand | 562-566 | 5 | 70 |
| α-helix | 580-582 | 3 | |
| α-helix | 588-593 | 6 | |
| β-strand | 596-600 | 5 | 70 |
| α-helix | 606-622 | 17 | |
| α-helix | 631-634 | 4 | |
| α-helix | 635-647 | 13 | |
| β-strand | 651 | 1 | 75 |
| α-helix | 654-674 | 21 | |
| α-helix | 686-702 | 17 | |
| β-strand | 707 | 1 | 75 |
| α-helix | 710-724 | 15 | |
| α-helix | 725-727 | 3 | |
Chain A: 8 helices, 5 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 4-22 | 19 | |
| α-helix | 29-31 | 3 | |
| α-helix | 35-59 | 25 | |
| α-helix | 65-103 | 39 | |
| α-helix | 120-122 | 3 | |
| α-helix | 125-142 | 18 | |
| α-helix | 157-158 | 2 | |
| β-strand | 162-167 | 6 | 1 |
| β-strand | 173-175 | 3 | 2 |
| β-strand | 180-182 | 3 | 2 |
| β-strand | 188-192 | 5 | 1 |
| α-helix | 197-202 | 6 | |
| β-strand | 205-206 | 2 | 1 |
Chain B: 8 helices, 4 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 13-20 | 8 | |
| β-strand | 24-29 | 6 | 3 |
| α-helix | 58-61 | 4 | |
| β-strand | 69-72 | 4 | 3 |
| α-helix | 73-82 | 10 | |
| β-strand | 85-87 | 3 | 3 |
| α-helix | 95-107 | 13 | |
| α-helix | 121-129 | 9 | |
| α-helix | 138-159 | 22 | |
| α-helix | 160-162 | 3 | |
| β-strand | 168-169 | 2 | 4 |
| α-helix | 175-199 | 25 | |
5 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 |
|---|
| DNA replication complex GINS protein PSF1 | A | protein | 208 | Saccharomyces cerevisiae W303 | Q12488 (AlphaFold model) |
| DNA replication complex GINS protein PSF2 | B | protein | 213 | Saccharomyces cerevisiae W303 | P40359 (AlphaFold model) |
| DNA replication complex GINS protein PSF3 | C | protein | 217 | Saccharomyces cerevisiae W303 | Q12146 (AlphaFold model) |
| DNA replication complex GINS protein SLD5 | D | protein | 294 | Saccharomyces cerevisiae W303 | Q03406 (AlphaFold model) |
| Cell division control protein 45 | E | protein | 650 | Saccharomyces cerevisiae W303 | Q08032 |
| Leading strand DNA template | F | DNA | 48 | synthetic construct | |
| Lagging strand DNA template | G | DNA | 20 | synthetic construct | |
| DNA replication licensing factor MCM2 | 2 | protein | 868 | Saccharomyces cerevisiae W303 | P29469 |
| DNA replication licensing factor MCM3 | 3 | protein | 971 | Saccharomyces cerevisiae W303 | P24279 |
| DNA replication licensing factor MCM4 | 4 | protein | 933 | Saccharomyces cerevisiae W303 | P30665 |
| Minichromosome maintenance protein 5 | 5 | protein | 775 | Saccharomyces cerevisiae W303 | P29496 |
| DNA replication licensing factor MCM6 | 6 | protein | 1017 | Saccharomyces cerevisiae W303 | P53091 |
3 more molecules are not listed.
Sequence of entity 1 (A), FASTA
>9E2X_1 DNA replication complex GINS protein PSF1 (chains A)
MYGDLGNKLVLEAKRTKQLYARSNQDVNLPMYHEDIIRNILKEVSNLRKNTEYLKEQQQL
GMLDDKVAKCQYFVTLLCMERNKRCLLAYQRLRTDILDSMAWNNNGLDLMSSITFSQQDT
NNLSHQEQEYLKEYCDLITDLKSGDLVDIDLSGSLVPPSDVFIDVRVLKDAGEIQTEYGV
FNLIKDSQFFVRQSDVERLIQQGYLQKI
Sequence of entity 2 (B), FASTA
>9E2X_2 DNA replication complex GINS protein PSF2 (chains B)
MSLPAHLQQTFSPEEIQFIVENEPIKIFPRITTRQKIRGDDRGTGNHTRWQLITTDDKAL
NNMVAMRSTEVVLWIALLLKQQSKCSIVAPQWLTTKELDRKIQYEKTHPDRFSELPWNWL
VLARILFNKAKDDFHDPIHELRGKIQDLREIRQIKVLKGLKYLNESHLQLDNLSLLEINE
LRPFITEIMDKLREIHTASLTAGTENDEEEFNI
Sequence of entity 3 (C), FASTA
>9E2X_3 DNA replication complex GINS protein PSF3 (chains C)
MGSSHHHHHHSSGLVPRGSHMASMGYYDIDDVLADGTEFPCKFQYDIPGLGYLENNPGRP
ITKNTKLSLPLWLARILAIVGGDEALVDEEPVPFVELLPPDMFSTKVMNAIKTDPVALDL
HSINSHFFSLAIKWIMLFSEKELANVVSELLLQRAQELNHHASSLSIDLNADSTGKNSAN
TNIATSTFLLKLEEMEKEIYKKSHESYKDTKRWMFKK
Sequence of entity 4 (D), FASTA
>9E2X_4 DNA replication complex GINS protein SLD5 (chains D)
MDINIDDILAELDKETTAVDSTKITQGSSSTTHRDANTIVGSSLDLNDKTQIYVSPQQDF
SDLMKSWKNERCSPELLPYPHQLMKRLLNRISMQSQLIENISMGFLDMQNASNANPPMPN
ESKLPLLCMETELERLKFVIRSYIRCRLSKIDKFSLYLRQLNEDENSLISLTDLLSKDEI
KYHDTHSLIWLKLVNDSILKYMPEELQAINDTEGSVNMIDEPDWNKFVFIHVNGPPDGKW
NEDPLLQENEFGKPCYTVTIPDLKEEVELTIGSIYVMRYEVIRDLLRDDKVALI
Sequence of entity 5 (E), FASTA
>9E2X_5 Cell division control protein 45 (chains E)
MYYGISQFSEAYNKILRNSSSHSSCQLVIFVSCLNIDALCATKMLSLLFKKQLVQSQIVP
IFGYSELRRHYSQLDDNINSLLLVGFGGVIDLEAFLEIDPQEYVIDTDEKSGEQSFRRDI
YVLDAHRPWNLDNIFGSQIIQCFDDGTVDDTLGEQKEAYYKLLELDEESGDDELSGDEND
NNGGDDEATDADEVTDEDEEDEDETISNKRGNSSIGPNDLSKRKQRKKQIHEYEGVLEEY
YSQGTTVVNSISAQIYSLLSAIGETNLSNLWLNILGTTSLDIAYAQVYNRLYPLLQDEVK
RLTPSSRNSVKTPDTLTLNIQPDYYLFLLRHSSLYDSFYYSNYVNAKLSLWNENGKKRLH
KMFARMGIPLSTAQETWLYMDHSIKRELGIIFDKNLDRYGLQDIIRDGFVRTLGYRGSIS
ASEFVEALTALLEVGNSTDKDSVKINNDNNDDTDGEEEEDNSAQKLTNLRKRWVSNFWLS
WDALDDRKVELLNRGIQLAQDLQRAIFNTGVAILEKKLIKHLRIYRLCVLQDGPDLDLYR
NPLTLLRLGNWLIECCAESEDKQLLPMVLASIDENTDTYLVAGLTPRYPRGLDTIHTKKP
ILNNFSMAFQQITAETDAKVRIDNFESSIIEIRREDLSPFLEKLTLSGLL
Sequence of entity 6 (F), FASTA
>9E2X_6 Leading strand DNA template (chains F)
TCGTGCTGAGTGATATCTGCTTTGGGTGGGTGGGTGGGTTGAGGCAAT
Sequence of entity 7 (G), FASTA
>9E2X_7 Lagging strand DNA template (chains G)
GCAGATATCACTCAGCACGA
Sequence of entity 8 (2), FASTA
>9E2X_8 DNA replication licensing factor MCM2 (chains 2)
MSDNRRRRREEDDSDSENELPPSSPQQHFRGGMNPVSSPIGSPDMINPEGDDNEVDDVPD
IDEVEEQMNEVDLMDDNMYEDYAADHNRDRYDPDQVDDREQQELSLSERRRIDAQLNERD
RLLRNVAYIDDEDEEQEGAAQLDEMGLPVQRRRRRRQYEDLENSDDDLLSDMDIDPLREE
LTLESLSNVKANSYSEWITQPNVSRTIARELKSFLLEYTDETGRSVYGARIRTLGEMNSE
SLEVNYRHLAESKAILALFLAKCPEEMLKIFDLVAMEATELHYPDYARIHSEIHVRISDF
PTIYSLRELRESNLSSLVRVTGVVTRRTGVFPQLKYVKFNCLKCGSILGPFFQDSNEEIR
ISFCTNCKSKGPFRVNGEKTVYRNYQRVTLQEAPGTVPPGRLPRHREVILLADLVDVSKP
GEEVEVTGIYKNNYDGNLNAKNGFPVFATIIEANSIKRREGNTANEGEEGLDVFSWTEEE
EREFRKISRDRGIIDKIISSMAPSIYGHRDIKTAVACSLFGGVPKNVNGKHSIRGDINVL
LLGDPGTAKSQILKYVEKTAHRAVFATGQGASAVGLTASVRKDPITKEWTLEGGALVLAD
KGVCLIDEFDKMNDQDRTSIHEAMEQQSISISKAGIVTTLQARCSIIAAANPNGGRYNST
LPLAQNVSLTEPILSRFDILCVVRDLVDEEADERLATFVVDSHVRSHPENDEDREGEELK
NNGESAIEQGEDEINEQLNARQRRLQRQRKKEEEISPIPQELLMKYIHYARTKIYPKLHQ
MDMDKVSRVYADLRRESISTGSFPITVRHLESILRIAESFAKMRLSEFVSSYDLDRAIKV
VVDSFVDAQKVSVRRQLRRSFAIYTLGH
Sequence of entity 9 (3), FASTA
>9E2X_9 DNA replication licensing factor MCM3 (chains 3)
MEGSTGFDGDATTFFAPDAVFGDRVRRFQEFLDTFTSYRDSVRSIQVYNSNNAANYNDDQ
DDADERDLLGDDDGDDLEKEKKAASSTSLNILPHRIIISLDDLREFDRSFWSGILVEPAY
FIPPAEKALTDLADSMDDVPHPNASAVSSRHPWKLSFKGSFGAHALSPRTLTAQHLNKLV
SVEGIVTKTSLVRPKLIRSVHYAAKTGRFHYRDYTDATTTLTTRIPTPAIYPTEDTEGNK
LTTEYGYSTFIDHQRITVQEMPEMAPAGQLPRSIDVILDDDLVDKTKPGDRVNVVGVFKS
LGAGGMNQSNSNTLIGFKTLILGNTVYPLHARSTGVAARQMLTDFDIRNINKLSKKKDIF
DILSQSLAPSIYGHDHIKKAILLMLMGGVEKNLENGSHLRGDINILMVGDPSTAKSQLLR
FVLNTASLAIATTGRGSSGVGLTAAVTTDRETGERRLEAGAMVLADRGVVCIDEFDKMTD
VDRVAIHEVMEQQTVTIAKAGIHTTLNARCSVIAAANPVFGQYDVNRDPHQNIALPDSLL
SRFDLLFVVTDDINEIRDRSISEHVLRTHRYLPPGYLEGEPVRERLNLSLAVGEDADINP
EEHSNSGAGVENEGEDDEDHVFEKFNPLLQAGAKLAKNKGNYNGTEIPKLVTIPFLRKYV
QYAKERVIPQLTQEAINVIVKNYTDLRNDDNTKKSPITARTLETLIRLATAHAKVRLSKT
VNKVDAKVAANLLRFALLGEDIGNDIDEEESEYEEALSKRSPQKSPKKRQRVRQPASNSG
SPIKSTPRRSTASSVNATPSSARRILRFQDDEQNAGEDDNDIMSPLPADEEAELQRRLQL
GLRVSPRRREHLHAPEEGSSGPLTEVGTPRLPNVSSAGQDDEQQQSVISFDNVEPGTIST
GRLSLISGIIARLMQTEIFEEESYPVASLFERINEELPEEEKFSAQEYLAGLKIMSDRNN
LMVADDKVWRV
Sequence of entity 10 (4), FASTA
>9E2X_10 DNA replication licensing factor MCM4 (chains 4)
MSQQSSSPTKEDNNSSSPVVPNPDSVPPQLSSPALFYSSSSSQGDIYGRNNSQNLSQGEG
NIRAAIGSSPLNFPSSSQRQNSDVFQSQGRQGRIRSSASASGRSRYHSDLRSDRALPTSS
SSLGRNGQNRVHMRRNDIHTSDLSSPRRIVDFDTRSGVNTLDTSSSSAPPSEASEPLRII
WGTNVSIQECTTNFRNFLMSFKYKFRKILDEREEFINNTTDEELYYIKQLNEMRELGTSN
LNLDARNLLAYKQTEDLYHQLLNYPQEVISIMDQTIKDCMVSLIVDNNLDYDLDEIETKF
YKVRPYNVGSCKGMRELNPNDIDKLINLKGLVLRSTPVIPDMKVAFFKCNVCDHTMAVEI
DRGVIQEPARCERIDCNEPNSMSLIHNRCSFADKQVIKLQETPDFVPDGQTPHSISLCVY
DELVDSCRAGDRIEVTGTFRSIPIRANSRQRVLKSLYKTYVDVVHVKKVSDKRLDVDTST
IEQELMQNKVDHNEVEEVRQITDQDLAKIREVAAREDLYSLLARSIAPSIYELEDVKKGI
LLQLFGGTNKTFTKGGRYRGDINILLCGDPSTSKSQILQYVHKITPRGVYTSGKGSSAVG
LTAYITRDVDTKQLVLESGALVLSDGGVCCIDEFDKMSDSTRSVLHEVMEQQTISIAKAG
IITTLNARSSILASANPIGSRYNPNLPVTENIDLPPPLLSRFDLVYLVLDKVDEKNDREL
AKHLTNLYLEDKPEHISQDDVLPVEFLTMYISYAKEHIHPIITEAAKTELVRAYVGMRKM
GDDSRSDEKRITATTRQLESMIRLAEAHAKMKLKNVVELEDVQEAVRLIRSAIKDYATDP
KTGKIDMNLVQTGKSVIQRKLQEDLSREIMNVLKDQASDSMSFNELIKQINEHSQDRVES
SDIQEALSRLQQEDKVIVLGEGVRRSVRLNNRV
Sequence of entity 11 (5), FASTA
>9E2X_11 Minichromosome maintenance protein 5 (chains 5)
MSFDRPEIYSAPVLQGESPNDDDNTEIIKSFKNFILEFRLDSQFIYRDQLRNNILVKNYS
LTVNMEHLIGYNEDIYKKLSDEPSDIIPLFETAITQVAKRISILSRAQSANNNDKDPENT
SMDTDSLLLNSLPTFQLILNSNANQIPLRDLDSEHVSKIVRLSGIIISTSVLSSRATYLS
IMCRNCRHTTSITINNFNSITGNTVSLPRSCLSTIESESSMANESNIGDESTKKNCGPDP
YIIIHESSKFIDQQFLKLQEIPELVPVGEMPRNLTMTCDRYLTNKVIPGTRVTIVGIYSI
YNSKNGAGSGRSGGGNGGSGVAIRTPYIKILGIQSDVETSSIWNSVTMFTEEEEEEFLQL
SRNPKLYEILTNSIAPSIFGNEDIKKAIVCLLMGGSKKILPDGMRLRGDINVLLLGDPGT
AKSQLLKFVEKVSPIAVYTSGKGSSAAGLTASVQRDPMTREFYLEGGAMVLADGGVVCID
EFDKMRDEDRVAIHEAMEQQTISIAKAGITTVLNSRTSVLAAANPIYGRYDDLKSPGDNI
DFQTTILSRFDMIFIVKDDHNEERDISIANHVINIHTGNANAMQNQQEENGSEISIEKMK
RYITYCRLKCAPRLSPQAAEKLSSNFVTIRKQLLINELESTERSSIPITIRQLEAIIRIT
ESLAKLELSPIAQERHVDEAIRLFQASTMDAASQDPIGGLNQASGTSLSEIRRFEQELKR
RLPIGWSTSYQTLRREFVDTHRFSQLALDKALYALEKHETIQLRHQGQNIYRSGV
Sequence of entity 12 (6), FASTA
>9E2X_12 DNA replication licensing factor MCM6 (chains 6)
MSSPFPADTPSSNRPSNSSPPPSSIGAGFGSSSGLDSQIGSRLHFPSSSQPHVSNSQTGP
FVNDSTQFSSQRLQTDGSATNDMEGNEPARSFKSRALNHVKKVDDVTGEKVREAFEQFLE
DFSVQSTDTGEVEKVYRAQIEFMKIYDLNTIYIDYQHLSMRENGALAMAISEQYYRFLPF
LQKGLRRVVRKYAPELLNTSDSLKRSEGDEGQADEDEQQDDDMNGSSLPRDSGSSAAPGN
GTSAMATRSITTSTSPEQTERVFQISFFNLPTVHRIRDIRSEKIGSLLSISGTVTRTSEV
RPELYKASFTCDMCRAIVDNVEQSFKYTEPTFCPNPSCENRAFWTLNVTRSRFLDWQKVR
IQENANEIPTGSMPRTLDVILRGDSVERAKPGDRCKFTGVEIVVPDVTQLGLPGVKPSST
LDTRGISKTTEGLNSGVTGLRSLGVRDLTYKISFLACHVISIGSNIGASSPDANSNNRET
ELQMAANLQANNVYQDNERDQEVFLNSLSSDEINELKEMVKDEHIYDKLVRSIAPAVFGH
EAVKKGILLQMLGGVHKSTVEGIKLRGDINICVVGDPSTSKSQFLKYVVGFAPRSVYTSG
KASSAAGLTAAVVRDEEGGDYTIEAGALMLADNGICCIDEFDKMDISDQVAIHEAMEQQT
ISIAKAGIHATLNARTSILAAANPVGGRYNRKLSLRGNLNMTAPIMSRFDLFFVILDDCN
EKIDTELASHIVDLHMKRDEAIEPPFSAEQLRRYIKYARTFKPILTKEARSYLVEKYKEL
RKDDAQGFSRSSYRITVRQLESMIRLSEAIARANCVDEITPSFIAEAYDLLRQSIIRVDV
DDVEMDEEFDNIESQSHAASGNNDDNDDGTGSGVITSEPPADIEEGQSEATARPGTSEKK
KTTVTYDKYVSMMNMIVRKIAEVDREGAEELTAVDIVDWYLLQKENDLGSLAEYWEERRL
AFKVIKRLVKDRILMEIHGTRHNLRDLENEENENNKTVYVIHPNCEVLDQLEPQDSS
Ligands and cofactors
| ID | Name | Formula | Copies |
|---|
| ATP | Adenosine-5'-triphosphate | C10 H16 N5 O13 P3 | 4 |
| MG | Magnesium ion | Mg | 5 |
| ZN | Zinc ion | Zn | 5 |
| ADP | Adenosine-5'-diphosphate | C10 H15 N5 O10 P2 | 2 |
Primary citation
G-quadruplex-stalled eukaryotic replisome structure reveals helical inchworm DNA translocation. Batra, S., Allwein, B., Kumar, C. et al. Science (2025) 387:eadt1978-eadt1978. DOI 10.1126/science.adt1978 · PubMed
Other PDB entries of the same protein (UniProt Q12488 (AlphaFold model), which also has an AlphaFold model), best resolution first:
- 28VY 2.7 Å, sCMGE assembled on ARS1 DNA with Sld2 and RPA
- 8KG6 3.07 Å, Yeast replisome in state I
- 7PMK 3.2 Å, S. cerevisiae replisome-SCF(Dia2) complex bound to double-stranded DNA (conformation I)
- 7PMN 3.2 Å, S. cerevisiae replisome-SCF(Dia2) complex bound to double-stranded DNA (conformation II)
- 9E2Y 3.2 Å, Cryo-EM structure of yeast CMG helicase stalled at G4-containing DNA template, state 3
- 9RHI 3.2 Å, pre-Initiation Complex on ARS1 DNA (monomer)
- 9E2W 3.3 Å, Cryo-EM structure of yeast CMG helicase stalled at G4-containing DNA template, state 1
- 9RHJ 3.4 Å, Pre-Initiation Complex on ARS1 DNA (dimer)
- 8B9A 3.5 Å, S. cerevisiae replisome + Ctf4, bound by pol alpha primase. Complex engaged with a fork…
- 8B9B 3.5 Å, S. cerevisiae replisome + Ctf4, bound by pol alpha. Complex engaged with a fork DNA…
- 3JC6 3.7 Å, Structure of the eukaryotic replicative CMG helicase and pumpjack motion
- 6SKL 3.7 Å, Cryo-EM structure of the CMG Fork Protection Complex at a replication fork -…
Browse structure collections
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