9E2W: DNA replication licensing factor MCM2
Cryo-EM structure of yeast CMG helicase stalled at G4-containing DNA template, state 1. Determined by electron microscopy at 3.3 Å resolution. Released 26 Mar 2025.
- Method
- Electron microscopy
- Resolution
- 3.3 Å
- Organisms
- Saccharomyces cerevisiae W303, synthetic construct
- Chains
- 15
- Atoms
- 48,591
- Mol. weight
- 965.6 kDa
- Ligands
- ATP, ZN, MG, ADP
- Released
- 26 Mar 2025
Explore 9E2W in 3D
Show helices and sheets
RCSB PDB
PDBe
Secondary structure: helices and β-sheets
9E2W contains 312 α-helices and 229 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: 33 helices, 31 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 | 1 |
| β-strand | 225 | 1 | 1 |
| α-helix | 226-236 | 11 | |
| β-strand | 241-244 | 4 | 2 |
| α-helix | 246-252 | 7 | |
| α-helix | 254-262 | 9 | |
| α-helix | 264-282 | 19 | |
| α-helix | 286-288 | 3 | |
| β-strand | 294-297 | 4 | 2 |
| β-strand | 304 | 1 | 3 |
| α-helix | 311-313 | 3 | |
| β-strand | 317-327 | 11 | 3 |
| β-strand | 331-334 | 4 | 3 |
| β-strand | 336-337 | 2 | 4 |
| β-strand | 339-341 | 3 | 5 |
| β-strand | 347 | 1 | 5 |
| α-helix | 348-350 | 3 | |
| β-strand | 351-352 | 2 | 4 |
| β-strand | 373-375 | 3 | 5 |
| β-strand | 382-391 | 10 | 3 |
| α-helix | 398 | 1 | |
| β-strand | 405-411 | 7 | 3 |
| β-strand | 423-433 | 11 | 3 |
| α-helix | 436-442 | 7 | |
| β-strand | 446 | 1 | 6 |
| β-strand | 448-458 | 11 | 3 |
| α-helix | 463-465 | 3 | |
| α-helix | 478-489 | 12 | |
| α-helix | 493-499 | 7 | |
| α-helix | 509-520 | 12 | |
| β-strand | 525-527 | 3 | 7 |
| β-strand | 531-533 | 3 | 7 |
| β-strand | 539-543 | 5 | 8 |
| α-helix | 550-559 | 10 | |
| β-strand | 563-567 | 5 | 8 |
| β-strand | 571 | 1 | 9 |
| β-strand | 578-582 | 5 | 10 |
| β-strand | 589-593 | 5 | 10 |
| α-helix | 595-598 | 4 | |
| β-strand | 603-607 | 5 | 8 |
| α-helix | 609-611 | 3 | |
| α-helix | 614-626 | 13 | |
| β-strand | 628-633 | 6 | 11 |
| β-strand | 636-641 | 6 | 11 |
| β-strand | 645-650 | 6 | 8 |
| α-helix | 653-655 | 3 | |
| α-helix | 663-666 | 4 | |
| α-helix | 671-675 | 5 | |
| β-strand | 679-683 | 5 | 8 |
| α-helix | 689-705 | 17 | |
| α-helix | 740-755 | 16 | |
| α-helix | 760-773 | 14 | |
| β-strand | 777 | 1 | 12 |
| α-helix | 783-797 | 15 | |
| α-helix | 803-804 | 2 | |
| α-helix | 807-823 | 17 | |
| β-strand | 828 | 1 | 12 |
| α-helix | 831-847 | 17 | |
| α-helix | 851-860 | 10 | |
| α-helix | 861-863 | 3 | |
Chain 3: 30 helices, 28 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 20-34 | 15 | |
| α-helix | 36-51 | 16 | |
| β-strand | 95-99 | 5 | 13 |
| 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 | 13 |
| β-strand | 165-166 | 2 | 14 |
| α-helix | 173-175 | 3 | |
| β-strand | 179-189 | 11 | 14 |
| α-helix | 190-192 | 3 | |
| β-strand | 193-203 | 11 | 14 |
| β-strand | 208-213 | 6 | 14 |
| α-helix | 230-232 | 3 | |
| β-strand | 234 | 1 | 15 |
| β-strand | 240 | 1 | 15 |
| β-strand | 242-243 | 2 | 14 |
| β-strand | 249-259 | 11 | 14 |
| α-helix | 262-264 | 3 | |
| α-helix | 270-272 | 3 | |
| β-strand | 273-279 | 7 | 14 |
| α-helix | 280-282 | 3 | |
| β-strand | 291-301 | 11 | 14 |
| β-strand | 318-328 | 11 | 14 |
| α-helix | 340-343 | 4 | |
| α-helix | 344-354 | 11 | |
| α-helix | 359-365 | 7 | |
| α-helix | 375-386 | 12 | |
| β-strand | 391-392 | 2 | 16 |
| β-strand | 398-399 | 2 | 16 |
| β-strand | 405-409 | 5 | 17 |
| α-helix | 415-425 | 11 | |
| β-strand | 429-433 | 5 | 17 |
| β-strand | 437 | 1 | 18 |
| α-helix | 439-443 | 5 | |
| β-strand | 444-448 | 5 | 19 |
| β-strand | 455-459 | 5 | 19 |
| α-helix | 461-464 | 4 | |
| β-strand | 469-473 | 5 | 17 |
| α-helix | 475-477 | 3 | |
| α-helix | 480-492 | 13 | |
| β-strand | 494-499 | 6 | 20 |
| β-strand | 502-507 | 6 | 20 |
| β-strand | 511-516 | 6 | 17 |
| α-helix | 529-532 | 4 | |
| α-helix | 537-541 | 5 | |
| β-strand | 545-549 | 5 | 17 |
| α-helix | 555-569 | 15 | |
| β-strand | 579 | 1 | 21 |
| α-helix | 580-582 | 3 | |
| α-helix | 650-652 | 3 | |
| α-helix | 653-666 | 14 | |
| β-strand | 670 | 1 | 22 |
| α-helix | 673-687 | 15 | |
| α-helix | 699-715 | 17 | |
| β-strand | 720 | 1 | 22 |
| α-helix | 723-738 | 16 | |
Chain 4: 28 helices, 34 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 341-344 | 4 | 23 |
| β-strand | 350 | 1 | 23 |
| α-helix | 351-363 | 13 | |
| β-strand | 366 | 1 | 24 |
| α-helix | 367-373 | 7 | |
| α-helix | 377-379 | 3 | |
| β-strand | 388 | 1 | 24 |
| α-helix | 390-400 | 11 | |
| β-strand | 404-408 | 5 | 25 |
| α-helix | 409-414 | 6 | |
| α-helix | 419-427 | 9 | |
| α-helix | 429-451 | 23 | |
| α-helix | 457-461 | 5 | |
| β-strand | 466-470 | 5 | 25 |
| α-helix | 483-485 | 3 | |
| β-strand | 489-499 | 11 | 26 |
| α-helix | 500-502 | 3 | |
| β-strand | 503-513 | 11 | 26 |
| β-strand | 519-523 | 5 | 26 |
| β-strand | 524-525 | 2 | 27 |
| β-strand | 528-529 | 2 | 27 |
| β-strand | 534 | 1 | 28 |
| β-strand | 543 | 1 | 28 |
| β-strand | 546-548 | 3 | 26 |
| α-helix | 550-552 | 3 | |
| β-strand | 554-565 | 12 | 26 |
| α-helix | 567-569 | 3 | |
| α-helix | 575-577 | 3 | |
| β-strand | 578-584 | 7 | 26 |
| α-helix | 585-587 | 3 | |
| β-strand | 596-606 | 11 | 26 |
| β-strand | 609 | 1 | 29 |
| β-strand | 617-618 | 2 | 29 |
| β-strand | 620 | 1 | 30 |
| α-helix | 621 | 1 | |
| β-strand | 622-632 | 11 | 26 |
| α-helix | 667-678 | 12 | |
| α-helix | 682-690 | 9 | |
| α-helix | 698-709 | 12 | |
| β-strand | 714-716 | 3 | 31 |
| β-strand | 720-722 | 3 | 31 |
| β-strand | 728-732 | 5 | 32 |
| α-helix | 739-748 | 10 | |
| β-strand | 754-755 | 2 | 32 |
| β-strand | 768-772 | 5 | 33 |
| β-strand | 777-781 | 5 | 33 |
| α-helix | 784-787 | 4 | |
| β-strand | 792-795 | 4 | 32 |
| α-helix | 803-815 | 13 | |
| β-strand | 817-820 | 4 | 34 |
| β-strand | 827-830 | 4 | 34 |
| β-strand | 834-839 | 6 | 32 |
| α-helix | 852-856 | 5 | |
| α-helix | 860-863 | 4 | |
| β-strand | 868-872 | 5 | 32 |
| α-helix | 878-890 | 13 | |
| α-helix | 908-921 | 14 | |
| β-strand | 925-926 | 2 | 35 |
| α-helix | 928-944 | 17 | |
| α-helix | 960-975 | 16 | |
| β-strand | 980-981 | 2 | 35 |
| α-helix | 983-999 | 17 | |
Chain 5: 34 helices, 25 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 23-37 | 15 | |
| α-helix | 45-55 | 11 | |
| β-strand | 60-64 | 5 | 36 |
| 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 | 36 |
| α-helix | 143-147 | 5 | |
| α-helix | 148-150 | 3 | |
| α-helix | 153-155 | 3 | |
| β-strand | 159-169 | 11 | 37 |
| α-helix | 170-172 | 3 | |
| β-strand | 173-183 | 11 | 37 |
| β-strand | 189-193 | 5 | 37 |
| β-strand | 241-259 | 19 | 37 |
| α-helix | 260-261 | 2 | |
| α-helix | 262-264 | 3 | |
| α-helix | 270-272 | 3 | |
| β-strand | 273-279 | 7 | 37 |
| α-helix | 280-282 | 3 | |
| β-strand | 291-301 | 11 | 37 |
| β-strand | 323 | 1 | 3 |
| β-strand | 325-335 | 11 | 37 |
| α-helix | 339-343 | 5 | |
| α-helix | 351-361 | 11 | |
| α-helix | 366-373 | 8 | |
| α-helix | 382-393 | 12 | |
| β-strand | 398-399 | 2 | 38 |
| β-strand | 405-406 | 2 | 38 |
| β-strand | 412-416 | 5 | 39 |
| α-helix | 422-432 | 11 | |
| β-strand | 436-440 | 5 | 39 |
| β-strand | 444 | 1 | 11 |
| α-helix | 446-449 | 4 | |
| β-strand | 453-455 | 3 | 40 |
| β-strand | 462-464 | 3 | 40 |
| α-helix | 468-471 | 4 | |
| β-strand | 476-480 | 5 | 39 |
| α-helix | 482-484 | 3 | |
| α-helix | 487-490 | 4 | |
| α-helix | 491-493 | 3 | |
| α-helix | 494-498 | 5 | |
| β-strand | 501-506 | 6 | 18 |
| β-strand | 509-514 | 6 | 18 |
| β-strand | 518-523 | 6 | 39 |
| α-helix | 536-539 | 4 | |
| α-helix | 544-549 | 6 | |
| β-strand | 552-556 | 5 | 39 |
| α-helix | 562-577 | 16 | |
| α-helix | 596-609 | 14 | |
| β-strand | 611 | 1 | 21 |
| β-strand | 613-614 | 2 | 41 |
| α-helix | 616-636 | 21 | |
| α-helix | 650-666 | 17 | |
| β-strand | 671-672 | 2 | 41 |
| α-helix | 674-684 | 11 | |
| α-helix | 685-689 | 5 | |
| α-helix | 690-693 | 4 | |
Chain 6: 35 helices, 34 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 183-186 | 4 | 42 |
| α-helix | 188-190 | 3 | |
| α-helix | 192-193 | 2 | |
| α-helix | 195-210 | 16 | |
| β-strand | 212 | 1 | 43 |
| β-strand | 223 | 1 | 43 |
| α-helix | 224-235 | 12 | |
| β-strand | 239-243 | 5 | 44 |
| α-helix | 244-249 | 6 | |
| α-helix | 250-253 | 4 | |
| α-helix | 254-261 | 8 | |
| α-helix | 263-281 | 19 | |
| α-helix | 283-285 | 3 | |
| β-strand | 287 | 1 | 45 |
| α-helix | 348 | 1 | |
| β-strand | 349 | 1 | 45 |
| α-helix | 350 | 1 | |
| β-strand | 353-357 | 5 | 44 |
| β-strand | 362-363 | 2 | 6 |
| α-helix | 365-367 | 3 | |
| α-helix | 370-372 | 3 | |
| β-strand | 376-386 | 11 | 6 |
| α-helix | 387-389 | 3 | |
| β-strand | 390-400 | 11 | 6 |
| β-strand | 406-411 | 6 | 6 |
| β-strand | 433-435 | 3 | 6 |
| β-strand | 441-451 | 11 | 6 |
| α-helix | 458 | 1 | |
| α-helix | 462-464 | 3 | |
| β-strand | 465-471 | 7 | 6 |
| α-helix | 472-474 | 3 | |
| β-strand | 483-494 | 12 | 6 |
| α-helix | 496-499 | 4 | |
| β-strand | 501 | 1 | 46 |
| β-strand | 504 | 1 | 46 |
| α-helix | 524-525 | 2 | |
| β-strand | 526 | 1 | 47 |
| β-strand | 527 | 1 | 27 |
| β-strand | 537 | 1 | 47 |
| β-strand | 539-550 | 12 | 6 |
| α-helix | 590-596 | 7 | |
| α-helix | 599-610 | 12 | |
| α-helix | 614-620 | 7 | |
| α-helix | 630-641 | 12 | |
| β-strand | 646-647 | 2 | 48 |
| β-strand | 653-654 | 2 | 48 |
| β-strand | 660-664 | 5 | 49 |
| α-helix | 670-678 | 9 | |
| β-strand | 684-687 | 4 | 49 |
| β-strand | 700-703 | 4 | 50 |
| β-strand | 710-713 | 4 | 50 |
| α-helix | 716-719 | 4 | |
| β-strand | 724-727 | 4 | 49 |
| α-helix | 730-732 | 3 | |
| α-helix | 735-746 | 12 | |
| β-strand | 749-754 | 6 | 9 |
| β-strand | 757-762 | 6 | 9 |
| β-strand | 766-771 | 6 | 49 |
| α-helix | 784-787 | 4 | |
| α-helix | 792-797 | 6 | |
| β-strand | 800-804 | 5 | 49 |
| α-helix | 810-826 | 17 | |
| α-helix | 827-830 | 4 | |
| α-helix | 837-849 | 13 | |
| β-strand | 853-854 | 2 | 51 |
| α-helix | 856-873 | 18 | |
| α-helix | 886-902 | 17 | |
| β-strand | 907-908 | 2 | 51 |
| α-helix | 910-922 | 13 | |
Chain 7: 31 helices, 26 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 7-8 | 2 | 14 |
| α-helix | 14-27 | 14 | |
| α-helix | 62-71 | 10 | |
| β-strand | 77-81 | 5 | 52 |
| α-helix | 82-95 | 14 | |
| α-helix | 102-108 | 7 | |
| α-helix | 110-123 | 14 | |
| α-helix | 125-128 | 4 | |
| α-helix | 134-136 | 3 | |
| α-helix | 138-155 | 18 | |
| α-helix | 194-197 | 4 | |
| β-strand | 201-205 | 5 | 52 |
| α-helix | 206-208 | 3 | |
| α-helix | 209-214 | 6 | |
| β-strand | 224-225 | 2 | 30 |
| α-helix | 232-234 | 3 | |
| β-strand | 238-248 | 11 | 30 |
| α-helix | 249-251 | 3 | |
| β-strand | 252-262 | 11 | 30 |
| β-strand | 268-272 | 5 | 30 |
| β-strand | 277-278 | 2 | 29 |
| α-helix | 279-281 | 3 | |
| β-strand | 283 | 1 | 53 |
| α-helix | 287-292 | 6 | |
| β-strand | 297 | 1 | 53 |
| β-strand | 298-316 | 19 | 30 |
| α-helix | 319-321 | 3 | |
| β-strand | 330-336 | 7 | 30 |
| α-helix | 337-339 | 3 | |
| β-strand | 348-358 | 11 | 30 |
| α-helix | 363-368 | 6 | |
| β-strand | 371 | 1 | 14 |
| β-strand | 373-383 | 11 | 30 |
| α-helix | 410-418 | 9 | |
| α-helix | 426-437 | 12 | |
| β-strand | 456-460 | 5 | 54 |
| β-strand | 462 | 1 | 55 |
| β-strand | 464 | 1 | 55 |
| α-helix | 466-476 | 11 | |
| β-strand | 481-484 | 4 | 54 |
| β-strand | 520-524 | 5 | 54 |
| α-helix | 531-543 | 13 | |
| β-strand | 545-547 | 3 | 56 |
| β-strand | 556-558 | 3 | 56 |
| β-strand | 562-567 | 6 | 54 |
| α-helix | 580-583 | 4 | |
| α-helix | 588-593 | 6 | |
| β-strand | 596-600 | 5 | 54 |
| α-helix | 606-622 | 17 | |
| α-helix | 631-634 | 4 | |
| α-helix | 635-647 | 13 | |
| β-strand | 651 | 1 | 57 |
| α-helix | 654-675 | 22 | |
| α-helix | 686-703 | 18 | |
| β-strand | 707 | 1 | 57 |
| α-helix | 708-709 | 2 | |
| α-helix | 710-729 | 20 | |
Chain A: 9 helices, 5 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 4-22 | 19 | |
| α-helix | 29-32 | 4 | |
| α-helix | 34-59 | 26 | |
| α-helix | 65-103 | 39 | |
| α-helix | 120-122 | 3 | |
| α-helix | 125-142 | 18 | |
| α-helix | 145-147 | 3 | |
| α-helix | 157-158 | 2 | |
| β-strand | 162-167 | 6 | 58 |
| β-strand | 173-175 | 3 | 59 |
| β-strand | 180-182 | 3 | 59 |
| β-strand | 188-192 | 5 | 58 |
| α-helix | 197-202 | 6 | |
| β-strand | 205-207 | 3 | 58 |
Chain B: 9 helices, 4 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 13-20 | 8 | |
| β-strand | 24-29 | 6 | 60 |
| α-helix | 58-61 | 4 | |
| β-strand | 69-72 | 4 | 60 |
| α-helix | 73-81 | 9 | |
| β-strand | 85-87 | 3 | 60 |
| α-helix | 91-93 | 3 | |
| α-helix | 95-107 | 13 | |
| α-helix | 119-129 | 11 | |
| α-helix | 138-159 | 22 | |
| α-helix | 160-162 | 3 | |
| β-strand | 169 | 1 | 61 |
| 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 licensing factor MCM2 | 2 | protein | 868 | Saccharomyces cerevisiae W303 | P29469 (AlphaFold model) |
| DNA replication licensing factor MCM3 | 3 | protein | 971 | Saccharomyces cerevisiae W303 | P24279 (AlphaFold model) |
| DNA replication licensing factor MCM4 | 4 | protein | 933 | Saccharomyces cerevisiae W303 | P30665 (AlphaFold model) |
| Minichromosome maintenance protein 5 | 5 | protein | 775 | Saccharomyces cerevisiae W303 | P29496 (AlphaFold model) |
| DNA replication licensing factor MCM6 | 6 | protein | 1017 | Saccharomyces cerevisiae W303 | P53091 |
| DNA replication licensing factor MCM7 | 7 | protein | 845 | Saccharomyces cerevisiae W303 | P38132 |
| DNA replication complex GINS protein PSF1 | A | protein | 208 | Saccharomyces cerevisiae W303 | Q12488 |
| DNA replication complex GINS protein PSF2 | B | protein | 213 | Saccharomyces cerevisiae W303 | P40359 |
| DNA replication complex GINS protein PSF3 | C | protein | 217 | Saccharomyces cerevisiae W303 | Q12146 |
| DNA replication complex GINS protein SLD5 | D | protein | 294 | Saccharomyces cerevisiae W303 | Q03406 |
| Cell division control protein 45 | E | protein | 650 | Saccharomyces cerevisiae W303 | Q08032 |
| Leading strand DNA template | F | DNA | 48 | synthetic construct | |
3 more molecules are not listed.
Sequence of entity 1 (2), FASTA
>9E2W_1 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 2 (3), FASTA
>9E2W_2 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 3 (4), FASTA
>9E2W_3 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 4 (5), FASTA
>9E2W_4 Minichromosome maintenance protein 5 (chains 5)
MSFDRPEIYSAPVLQGESPNDDDNTEIIKSFKNFILEFRLDSQFIYRDQLRNNILVKNYS
LTVNMEHLIGYNEDIYKKLSDEPSDIIPLFETAITQVAKRISILSRAQSANNNDKDPENT
SMDTDSLLLNSLPTFQLILNSNANQIPLRDLDSEHVSKIVRLSGIIISTSVLSSRATYLS
IMCRNCRHTTSITINNFNSITGNTVSLPRSCLSTIESESSMANESNIGDESTKKNCGPDP
YIIIHESSKFIDQQFLKLQEIPELVPVGEMPRNLTMTCDRYLTNKVIPGTRVTIVGIYSI
YNSKNGAGSGRSGGGNGGSGVAIRTPYIKILGIQSDVETSSIWNSVTMFTEEEEEEFLQL
SRNPKLYEILTNSIAPSIFGNEDIKKAIVCLLMGGSKKILPDGMRLRGDINVLLLGDPGT
AKSQLLKFVEKVSPIAVYTSGKGSSAAGLTASVQRDPMTREFYLEGGAMVLADGGVVCID
EFDKMRDEDRVAIHEAMEQQTISIAKAGITTVLNSRTSVLAAANPIYGRYDDLKSPGDNI
DFQTTILSRFDMIFIVKDDHNEERDISIANHVINIHTGNANAMQNQQEENGSEISIEKMK
RYITYCRLKCAPRLSPQAAEKLSSNFVTIRKQLLINELESTERSSIPITIRQLEAIIRIT
ESLAKLELSPIAQERHVDEAIRLFQASTMDAASQDPIGGLNQASGTSLSEIRRFEQELKR
RLPIGWSTSYQTLRREFVDTHRFSQLALDKALYALEKHETIQLRHQGQNIYRSGV
Sequence of entity 5 (6), FASTA
>9E2W_5 DNA replication licensing factor MCM6 (chains 6)
MSSPFPADTPSSNRPSNSSPPPSSIGAGFGSSSGLDSQIGSRLHFPSSSQPHVSNSQTGP
FVNDSTQFSSQRLQTDGSATNDMEGNEPARSFKSRALNHVKKVDDVTGEKVREAFEQFLE
DFSVQSTDTGEVEKVYRAQIEFMKIYDLNTIYIDYQHLSMRENGALAMAISEQYYRFLPF
LQKGLRRVVRKYAPELLNTSDSLKRSEGDEGQADEDEQQDDDMNGSSLPRDSGSSAAPGN
GTSAMATRSITTSTSPEQTERVFQISFFNLPTVHRIRDIRSEKIGSLLSISGTVTRTSEV
RPELYKASFTCDMCRAIVDNVEQSFKYTEPTFCPNPSCENRAFWTLNVTRSRFLDWQKVR
IQENANEIPTGSMPRTLDVILRGDSVERAKPGDRCKFTGVEIVVPDVTQLGLPGVKPSST
LDTRGISKTTEGLNSGVTGLRSLGVRDLTYKISFLACHVISIGSNIGASSPDANSNNRET
ELQMAANLQANNVYQDNERDQEVFLNSLSSDEINELKEMVKDEHIYDKLVRSIAPAVFGH
EAVKKGILLQMLGGVHKSTVEGIKLRGDINICVVGDPSTSKSQFLKYVVGFAPRSVYTSG
KASSAAGLTAAVVRDEEGGDYTIEAGALMLADNGICCIDEFDKMDISDQVAIHEAMEQQT
ISIAKAGIHATLNARTSILAAANPVGGRYNRKLSLRGNLNMTAPIMSRFDLFFVILDDCN
EKIDTELASHIVDLHMKRDEAIEPPFSAEQLRRYIKYARTFKPILTKEARSYLVEKYKEL
RKDDAQGFSRSSYRITVRQLESMIRLSEAIARANCVDEITPSFIAEAYDLLRQSIIRVDV
DDVEMDEEFDNIESQSHAASGNNDDNDDGTGSGVITSEPPADIEEGQSEATARPGTSEKK
KTTVTYDKYVSMMNMIVRKIAEVDREGAEELTAVDIVDWYLLQKENDLGSLAEYWEERRL
AFKVIKRLVKDRILMEIHGTRHNLRDLENEENENNKTVYVIHPNCEVLDQLEPQDSS
Sequence of entity 6 (7), FASTA
>9E2W_6 DNA replication licensing factor MCM7 (chains 7)
MSAALPSIQLPVDYNNLFNEITDFLVTFKQDTLSSDATRNENEDENLDAENIEQHLLEKG
PKYMAMLQKVANRELNSVIIDLDDILQYQNEKFLQGTQADDLVSAIQQNANHFTELFCRA
IDNNMPLPTKEIDYKDDVLDVILNQRRLRNERMLSDRTNEIRSENLMDTTMDPPSSMNDA
LREVVEDETELFPPNLTRRYFLYFKPLSQNCARRYRKKAISSKPLSVRQIKGDFLGQLIT
VRGIITRVSDVKPAVEVIAYTCDQCGYEVFQEVNSRTFTPLSECTSEECSQNQTKGQLFM
STRASKFSAFQECKIQELSQQVPVGHIPRSLNIHVNGTLVRSLSPGDIVDVTGIFLPAPY
TGFKALKAGLLTETYLEAQFVRQHKKKFASFSLTSDVEERVMELITSGDVYNRLAKSIAP
EIYGNLDVKKALLLLLVGGVDKRVGDGMKIRGDINVCLMGDPGVAKSQLLKAICKISPRG
VYTTGKGSSGVGLTAAVMKDPVTDEMILEGGALVLADNGICCIDEFDKMDESDRTAIHEV
MEQQTISISKAGINTTLNARTSILAAANPLYGRYNPRLSPLDNINLPAALLSRFDILFLM
LDIPSRDDDEKLAEHVTYVHMHNKQPDLDFTPVEPSKMREYIAYAKTKRPVMSEAVNDYV
VQAYIRLRQDSKREMDSKFSFGQATPRTLLGIIRLSQALAKLRLADMVDIDDVEEALRLV
RVSKESLYQETNKSKEDESPTTKIFTIIKKMLQETGKNTLSYENIVKTVRLRGFTMLQLS
NCIQEYSYLNVWHLINEGNTLKFVDDGTMDTDQEDSLVSTPKLAPQTTASANVSAQDSDI
DLQDA
Sequence of entity 7 (A), FASTA
>9E2W_7 DNA replication complex GINS protein PSF1 (chains A)
MYGDLGNKLVLEAKRTKQLYARSNQDVNLPMYHEDIIRNILKEVSNLRKNTEYLKEQQQL
GMLDDKVAKCQYFVTLLCMERNKRCLLAYQRLRTDILDSMAWNNNGLDLMSSITFSQQDT
NNLSHQEQEYLKEYCDLITDLKSGDLVDIDLSGSLVPPSDVFIDVRVLKDAGEIQTEYGV
FNLIKDSQFFVRQSDVERLIQQGYLQKI
Sequence of entity 8 (B), FASTA
>9E2W_8 DNA replication complex GINS protein PSF2 (chains B)
MSLPAHLQQTFSPEEIQFIVENEPIKIFPRITTRQKIRGDDRGTGNHTRWQLITTDDKAL
NNMVAMRSTEVVLWIALLLKQQSKCSIVAPQWLTTKELDRKIQYEKTHPDRFSELPWNWL
VLARILFNKAKDDFHDPIHELRGKIQDLREIRQIKVLKGLKYLNESHLQLDNLSLLEINE
LRPFITEIMDKLREIHTASLTAGTENDEEEFNI
Sequence of entity 9 (C), FASTA
>9E2W_9 DNA replication complex GINS protein PSF3 (chains C)
MGSSHHHHHHSSGLVPRGSHMASMGYYDIDDVLADGTEFPCKFQYDIPGLGYLENNPGRP
ITKNTKLSLPLWLARILAIVGGDEALVDEEPVPFVELLPPDMFSTKVMNAIKTDPVALDL
HSINSHFFSLAIKWIMLFSEKELANVVSELLLQRAQELNHHASSLSIDLNADSTGKNSAN
TNIATSTFLLKLEEMEKEIYKKSHESYKDTKRWMFKK
Sequence of entity 10 (D), FASTA
>9E2W_10 DNA replication complex GINS protein SLD5 (chains D)
MDINIDDILAELDKETTAVDSTKITQGSSSTTHRDANTIVGSSLDLNDKTQIYVSPQQDF
SDLMKSWKNERCSPELLPYPHQLMKRLLNRISMQSQLIENISMGFLDMQNASNANPPMPN
ESKLPLLCMETELERLKFVIRSYIRCRLSKIDKFSLYLRQLNEDENSLISLTDLLSKDEI
KYHDTHSLIWLKLVNDSILKYMPEELQAINDTEGSVNMIDEPDWNKFVFIHVNGPPDGKW
NEDPLLQENEFGKPCYTVTIPDLKEEVELTIGSIYVMRYEVIRDLLRDDKVALI
Sequence of entity 11 (E), FASTA
>9E2W_11 Cell division control protein 45 (chains E)
MYYGISQFSEAYNKILRNSSSHSSCQLVIFVSCLNIDALCATKMLSLLFKKQLVQSQIVP
IFGYSELRRHYSQLDDNINSLLLVGFGGVIDLEAFLEIDPQEYVIDTDEKSGEQSFRRDI
YVLDAHRPWNLDNIFGSQIIQCFDDGTVDDTLGEQKEAYYKLLELDEESGDDELSGDEND
NNGGDDEATDADEVTDEDEEDEDETISNKRGNSSIGPNDLSKRKQRKKQIHEYEGVLEEY
YSQGTTVVNSISAQIYSLLSAIGETNLSNLWLNILGTTSLDIAYAQVYNRLYPLLQDEVK
RLTPSSRNSVKTPDTLTLNIQPDYYLFLLRHSSLYDSFYYSNYVNAKLSLWNENGKKRLH
KMFARMGIPLSTAQETWLYMDHSIKRELGIIFDKNLDRYGLQDIIRDGFVRTLGYRGSIS
ASEFVEALTALLEVGNSTDKDSVKINNDNNDDTDGEEEEDNSAQKLTNLRKRWVSNFWLS
WDALDDRKVELLNRGIQLAQDLQRAIFNTGVAILEKKLIKHLRIYRLCVLQDGPDLDLYR
NPLTLLRLGNWLIECCAESEDKQLLPMVLASIDENTDTYLVAGLTPRYPRGLDTIHTKKP
ILNNFSMAFQQITAETDAKVRIDNFESSIIEIRREDLSPFLEKLTLSGLL
Sequence of entity 12 (F), FASTA
>9E2W_12 Leading strand DNA template (chains F)
TCGTGCTGAGTGATATCTGCTTTGGGTGGGTGGGTGGGTTGAGGCAAT
Ligands and cofactors
| ID | Name | Formula | Copies |
|---|
| ATP | Adenosine-5'-triphosphate | C10 H16 N5 O13 P3 | 5 |
| ZN | Zinc ion | Zn | 5 |
| MG | Magnesium ion | Mg | 6 |
| ADP | Adenosine-5'-diphosphate | C10 H15 N5 O10 P2 | 1 |
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 P29469 (AlphaFold model), which also has an AlphaFold model), best resolution first:
- 7W8G 2.52 Å, Cryo-EM structure of MCM double hexamer
- 28VY 2.7 Å, sCMGE assembled on ARS1 DNA with Sld2 and RPA
- 8RIF 2.79 Å, Cryo-EM structure of the MCM double hexamer loaded onto dsDNA.
- 7V3V 2.9 Å, Cryo-EM structure of MCM double hexamer bound with DDK in State I
- 9TH8 2.9 Å, Cryo-EM structure of MCM2-7 DH bound to Sld3-Sld7, Cdc45 and DNA
- 7P30 3.0 Å, 3.0 A resolution structure of a DNA-loaded MCM double hexamer
- 8KG6 3.07 Å, Yeast replisome in state I
- 9RHL 3.1 Å, Phospho-DH bound by Sld3-MBD on ARS1 DNA
- 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)
- 7PT6 3.2 Å, Structure of MCM2-7 DH complexed with Cdc7-Dbf4 in the presence of ATPgS, state III
- 7V3U 3.2 Å, Cryo-EM structure of MCM double hexamer with structured Mcm4-NSD
Browse structure collections
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