9EEB: Tubulin cofactors D,E,G
Tubulin cofactors D,E,G bound to tubulin dimer. Determined by electron microscopy at 3.7 Å resolution. Released 4 Feb 2026.
- Method
- Electron microscopy
- Resolution
- 3.7 Å
- Organisms
- Saccharomyces cerevisiae, Sus scrofa
- Chains
- 6
- Atoms
- 12,531
- Mol. weight
- 291.52 kDa
- Ligands
- GTP, GDP
- Released
- 4 Feb 2026
Explore 9EEB in 3D
Show helices and sheets
RCSB PDB
PDBe
Secondary structure: helices and β-sheets
9EEB contains 151 α-helices and 70 β-strands across 6 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: 24 helices, 18 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 5-8 | 4 | 15 |
| α-helix | 11-28 | 18 | |
| β-strand | 35 | 1 | 16 |
| α-helix | 48-51 | 4 | |
| β-strand | 53-55 | 3 | 17 |
| β-strand | 60 | 1 | 16 |
| β-strand | 61-63 | 3 | 17 |
| β-strand | 65-69 | 5 | 15 |
| α-helix | 72-80 | 9 | |
| α-helix | 89-91 | 3 | |
| β-strand | 92-94 | 3 | 15 |
| α-helix | 103-104 | 2 | |
| α-helix | 105-110 | 6 | |
| α-helix | 111-127 | 17 | |
| β-strand | 134-140 | 7 | 15 |
| α-helix | 144 | 1 | |
| α-helix | 145-149 | 5 | |
| α-helix | 150-160 | 11 | |
| β-strand | 165-172 | 8 | 15 |
| α-helix | 173 | 1 | |
| α-helix | 183-195 | 13 | |
| β-strand | 200-205 | 6 | 15 |
| α-helix | 206-217 | 12 | |
| α-helix | 224-243 | 20 | |
| α-helix | 252-259 | 8 | |
| β-strand | 269-273 | 5 | 18 |
| β-strand | 277 | 1 | 19 |
| α-helix | 280-282 | 3 | |
| α-helix | 288-296 | 9 | |
| α-helix | 298-300 | 3 | |
| β-strand | 301 | 1 | 18 |
| β-strand | 312-321 | 10 | 18 |
| α-helix | 325-338 | 14 | |
| β-strand | 343 | 1 | 18 |
| β-strand | 351-356 | 6 | 18 |
| α-helix | 359-361 | 3 | |
| β-strand | 368 | 1 | 19 |
| β-strand | 373-381 | 9 | 18 |
| α-helix | 382-384 | 3 | |
| α-helix | 385-401 | 17 | |
| α-helix | 405-411 | 7 | |
| α-helix | 415-435 | 21 | |
Chain B: 27 helices, 18 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 3-9 | 7 | 8 |
| α-helix | 11-27 | 17 | |
| β-strand | 30 | 1 | 9 |
| β-strand | 35 | 1 | 10 |
| β-strand | 36 | 1 | 9 |
| α-helix | 41-45 | 5 | |
| α-helix | 47-49 | 3 | |
| β-strand | 51-54 | 4 | 10 |
| β-strand | 58-61 | 4 | 10 |
| β-strand | 63-67 | 5 | 8 |
| α-helix | 70-77 | 8 | |
| α-helix | 82-84 | 3 | |
| α-helix | 87-89 | 3 | |
| β-strand | 90-92 | 3 | 8 |
| α-helix | 101-102 | 2 | |
| α-helix | 103-107 | 5 | |
| α-helix | 108-125 | 18 | |
| β-strand | 130-138 | 9 | 8 |
| α-helix | 143-147 | 5 | |
| α-helix | 148-158 | 11 | |
| β-strand | 164-170 | 7 | 8 |
| α-helix | 171 | 1 | |
| α-helix | 181-195 | 15 | |
| β-strand | 199-203 | 5 | 8 |
| α-helix | 204-209 | 6 | |
| α-helix | 210-214 | 5 | |
| α-helix | 222-241 | 20 | |
| α-helix | 250-257 | 8 | |
| β-strand | 265-266 | 2 | 8 |
| β-strand | 267-271 | 5 | 11 |
| α-helix | 286-294 | 9 | |
| α-helix | 296-298 | 3 | |
| β-strand | 299 | 1 | 11 |
| α-helix | 305-307 | 3 | |
| β-strand | 310-318 | 9 | 11 |
| α-helix | 323-336 | 14 | |
| α-helix | 338-340 | 3 | |
| β-strand | 341 | 1 | 11 |
| β-strand | 349-354 | 6 | 11 |
| α-helix | 357-358 | 2 | |
| β-strand | 364-371 | 8 | 11 |
| α-helix | 372-374 | 3 | |
| α-helix | 375-391 | 17 | |
| α-helix | 395-401 | 7 | |
| α-helix | 405-426 | 22 | |
Chain D: 71 helices, 6 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 5-16 | 12 | |
| α-helix | 20-32 | 13 | |
| α-helix | 33-35 | 3 | |
| α-helix | 37-41 | 5 | |
| α-helix | 44-53 | 10 | |
| α-helix | 60-84 | 25 | |
| α-helix | 90-101 | 12 | |
| α-helix | 108-124 | 17 | |
| α-helix | 132-144 | 13 | |
| α-helix | 145-147 | 3 | |
| α-helix | 152-166 | 15 | |
| α-helix | 171-173 | 3 | |
| α-helix | 176-179 | 4 | |
| α-helix | 182-194 | 13 | |
| α-helix | 196-202 | 7 | |
| α-helix | 205-217 | 13 | |
| α-helix | 221-223 | 3 | |
| α-helix | 224-234 | 11 | |
| α-helix | 237-249 | 13 | |
| α-helix | 256-276 | 21 | |
| α-helix | 278-297 | 20 | |
| α-helix | 305-320 | 16 | |
| α-helix | 326-331 | 6 | |
| α-helix | 332-336 | 5 | |
| α-helix | 337-341 | 5 | |
| β-strand | 345-347 | 3 | 12 |
| β-strand | 350-352 | 3 | 12 |
| α-helix | 354-370 | 17 | |
| α-helix | 378-393 | 16 | |
| α-helix | 398-414 | 17 | |
| α-helix | 416-419 | 4 | |
| α-helix | 423-431 | 9 | |
| α-helix | 437-452 | 16 | |
| α-helix | 459-470 | 12 | |
| α-helix | 471-479 | 9 | |
| α-helix | 483-488 | 6 | |
| α-helix | 489-494 | 6 | |
| α-helix | 495-499 | 5 | |
| α-helix | 505-514 | 10 | |
| α-helix | 519-534 | 16 | |
| α-helix | 542-548 | 7 | |
| α-helix | 549-553 | 5 | |
| α-helix | 571-586 | 16 | |
| α-helix | 593-603 | 11 | |
| α-helix | 610-625 | 16 | |
| α-helix | 631-639 | 9 | |
| α-helix | 643-648 | 6 | |
| α-helix | 649-651 | 3 | |
| α-helix | 655-668 | 14 | |
| α-helix | 671-684 | 14 | |
| α-helix | 689-691 | 3 | |
| α-helix | 694-707 | 14 | |
| β-strand | 713-715 | 3 | 13 |
| β-strand | 717-718 | 2 | 13 |
| α-helix | 720-733 | 14 | |
| α-helix | 736-740 | 5 | |
| α-helix | 742-754 | 13 | |
| α-helix | 758-769 | 12 | |
| α-helix | 780-790 | 11 | |
| α-helix | 795-804 | 10 | |
| α-helix | 811-827 | 17 | |
| α-helix | 830-842 | 13 | |
| α-helix | 845-846 | 2 | |
| α-helix | 847-853 | 7 | |
| α-helix | 856-859 | 4 | |
| β-strand | 860 | 1 | 14 |
| α-helix | 866 | 1 | |
| β-strand | 867 | 1 | 14 |
| α-helix | 868-870 | 3 | |
| α-helix | 871-887 | 17 | |
| α-helix | 899-910 | 12 | |
| α-helix | 915-935 | 21 | |
| α-helix | 941-964 | 24 | |
| α-helix | 970-986 | 17 | |
| α-helix | 989-1000 | 12 | |
| α-helix | 1003-1006 | 4 | |
| α-helix | 1009-1012 | 4 | |
Chain E: 13 helices, 20 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 127-128 | 2 | 1 |
| α-helix | 141-150 | 10 | |
| β-strand | 156-158 | 3 | 1 |
| β-strand | 166 | 1 | 2 |
| α-helix | 167-175 | 9 | |
| β-strand | 182-184 | 3 | 1 |
| β-strand | 192 | 1 | 2 |
| α-helix | 197-199 | 3 | |
| β-strand | 207-209 | 3 | 1 |
| α-helix | 217-224 | 8 | |
| β-strand | 232-234 | 3 | 1 |
| β-strand | 241 | 1 | 3 |
| α-helix | 242-246 | 5 | |
| α-helix | 249-251 | 3 | |
| β-strand | 258-260 | 3 | 1 |
| β-strand | 265 | 1 | 3 |
| β-strand | 280-282 | 3 | 1 |
| β-strand | 301-303 | 3 | 1 |
| α-helix | 313-321 | 9 | |
| β-strand | 327-328 | 2 | 1 |
| α-helix | 348-355 | 8 | |
| α-helix | 371-385 | 15 | |
| α-helix | 396-404 | 9 | |
| β-strand | 426-431 | 6 | 4 |
| α-helix | 436-438 | 3 | |
| β-strand | 440-444 | 5 | 4 |
| β-strand | 449 | 1 | 5 |
| α-helix | 450-461 | 12 | |
| β-strand | 469-473 | 5 | 4 |
| β-strand | 479-481 | 3 | 4 |
| β-strand | 488 | 1 | 5 |
| α-helix | 489-492 | 4 | |
| β-strand | 499-503 | 5 | 4 |
Chain F: 8 helices, 0 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 14-24 | 11 | |
| α-helix | 27-36 | 10 | |
| α-helix | 50-56 | 7 | |
| α-helix | 60-69 | 10 | |
| α-helix | 83-90 | 8 | |
| α-helix | 93-101 | 9 | |
| α-helix | 116-122 | 7 | |
| α-helix | 126-135 | 10 | |
Chain G: 8 helices, 8 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 3-14 | 12 | |
| β-strand | 16-23 | 8 | 6 |
| α-helix | 29-36 | 8 | |
| β-strand | 53-60 | 8 | 6 |
| β-strand | 63-70 | 8 | 6 |
| α-helix | 77-79 | 3 | |
| α-helix | 80-83 | 4 | |
| β-strand | 89-95 | 7 | 6 |
| α-helix | 101-115 | 15 | |
| β-strand | 120 | 1 | 7 |
| β-strand | 124 | 1 | 7 |
| β-strand | 125-131 | 7 | 6 |
| α-helix | 133-135 | 3 | |
| α-helix | 139-156 | 18 | |
| β-strand | 163-168 | 6 | 6 |
| α-helix | 176-190 | 15 | |
Molecules and chains
| Molecule | Chains | Type | Length | Organism | UniProt |
|---|
| Protein PAC2 | E | protein | 347 | Saccharomyces cerevisiae | P39937 (AlphaFold model) |
| GTP-binding protein CIN4 | G | protein | 191 | Saccharomyces cerevisiae | P39110 (AlphaFold model) |
| Tubulin beta chain | B | protein | 445 | Sus scrofa | P02554 (AlphaFold model) |
| Designed ankyrin repeat protein -- darpin | F | protein | 169 | Saccharomyces cerevisiae | |
| Chromosome instability protein 1 | D | protein | 1020 | Saccharomyces cerevisiae | P40987 (AlphaFold model) |
| Tubulin alpha-1A chain | A | protein | 451 | Sus scrofa | P02550 |
Sequence of entity 1 (E), FASTA
>9EEB_1 Protein PAC2 (chains E)
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXSLVLKIKIRAGKKPSSDLDYWVLPSFTVRYVKSVICRKLNFDILNVKLF
HENSEGMINEIKYNFRPISDFNVVNGDIIHVSSPVNNKSIQKVNSPS
Sequence of entity 2 (G), FASTA
>9EEB_2 GTP-binding protein CIN4 (chains G)
MGLLSIIRKQKLRDKEIRCLILGLDNSGKSTIVNKLLPKDEQNNDGIMPTVGFQIHSLMI
KDVTISLWDIGGQRTLRPFWDNYFDKTQAMIWCIDVSLSMRFDETLQELKELINRDENRI
GYECAVIVVLNKIDLVEDKSELHRRCLLVESELKCLFKPDIRIELVKCSGVTGEGIDNLR
DRLVESCHFTQ
Sequence of entity 3 (B), FASTA
>9EEB_3 Tubulin beta chain (chains B)
MREIVHIQAGQCGNQIGAKFWEVISDEHGIDPTGSYHGDSDLQLERINVYYNEAAGNKYV
PRAILVDLEPGTMDSVRSGPFGQIFRPDNFVFGQSGAGNNWAKGHYTEGAELVDSVLDVV
RKESESCDCLQGFQLTHSLGGGTGSGMGTLLISKIREEYPDRIMNTFSVVPSPKVSDTVV
EPYNATLSVHQLVENTDETYCIDNEALYDICFRTLKLTTPTYGDLNHLVSATMSGVTTCL
RFPGQLNADLRKLAVNMVPFPRLHFFMPGFAPLTSRGSQQYRALTVPELTQQMFDAKNMM
AACDPRHGRYLTVAAVFRGRMSMKEVDEQMLNVQNKNSSYFVEWIPNNVKTAVCDIPPRG
LKMSATFIGNSTAIQELFKRISEQFTAMFRRKAFLHWYTGEGMDEMEFTEAESNMNDLVS
EYQQYQDATADEQGEFEEEGEEDEA
Sequence of entity 4 (F), FASTA
>9EEB_4 DESIGNED ANKYRIN REPEAT PROTEIN -- DARPin (chains F)
MRGSHHHHHHGSDLGKKLLEAARAGQDDEVRVLMANGADVNATDASGLTPLHLAATYGHL
EIVEVLLKHGADVSASDLMGSTPLHLAALIGHLEIVEVLLKHGADVNAVDTWGDTPLRLA
AVMGHLKIVEALLKHGADVNAQDKFGKTAYDTSIDNGSEDLAEILQKLN
Sequence of entity 5 (D), FASTA
>9EEB_5 Chromosome instability protein 1 (chains D)
MHHHHHHNNIRALLDSIQSGVQTVSPEKHQQTIAAINKFQDDPALLDTILPRCVPLLTKS
FFCMSQRDQKLVAELFYNLDKISHSKVLKSLDTSIFRLNEILNYLQDRASPSSFSDVLCV
YLNLSWLSVILLSPYAFKDKFNKTLQVSSRFENYPICIPPINKIKAVLYFKNFTDAFDQL
PEREQANVPFLNQFLKLFIQSSEKANYYFSNENLRHLQQVALSNDGIKLLPKLFQISFNH
GSHDILDAIVEFFHDHLSSNSTDTRFQLAHSFAKIAKFLHQADPASFIELIDYTIENTVS
LLQAPCDSIDSNELHTSLLIIAEVALAKILPIDLVDRVLTLIIPKTCHFQKSHFQIIKGH
HIRDSTNFIIWSVIRSNRSNSLSPQVLQSLLSHLLINAFFDPELIIRYSSFAALQELLGR
SNKSLALNQNDIALILQANWKDLPRSFEENSGLIRRLFNPENTSKSAVCVWKVFRDWSFN
WNLLENLHLTTMKLNIDYNLVPLIKSKLSSPALLQEVLNKAGSSVTQNCQILYLYLKLFE
NDVNCPKISEICIDIYQKKIKFQLTTQAKRQFNDNSPELFQIFVILKYWQLTGQNDFNQE
LFWKFVDIVSPQKKLNLYNEFIPIIQQIISQCVSLNYTRIVQLIKSDNELTCRSICHMPD
QEKMCSLFFSQFPLLSPQSRSLLIGELDHHWDVRISLLPSNSYRKFRNIIINCLDDYTIT
QQGDVGRLVRIQALKLMQSHPDFLSGDCDSINPKLTRLLAEPVPEIRKLSYQLLASATSQ
ITVLSDSSILNFRHKQGLSEEFWKGYAVSAGAIHFTDSQLTSSIDSFIVYYRSLSPSQQL
ELCHDLIRIIPSAKQIAESRIRDRNKDPLTGGMRFDTIKFTIHCVKFWTRIMESGLVVLH
PNFNFQGVFAKFYNLHLLDCTTLRVSVIKFFPFLAISCYHTMRENADQKNLSNIILKRLL
VLVKREYAATKSKFMTDQNVALQGMFQIFLELGVTRQLQALQVACQKHELANILESDITL
Sequence of entity 6 (A), FASTA
>9EEB_6 Tubulin alpha-1A chain (chains A)
MRECISIHVGQAGVQIGNACWELYCLEHGIQPDGQMPSDKTIGGGDDSFNTFFSETGAGK
HVPRAVFVDLEPTVIDEVRTGTYRQLFHPEQLITGKEDAANNYARGHYTIGKEIIDLVLD
RIRKLADQCTGLQGFSVFHSFGGGTGSGFTSLLMERLSVDYGKKSKLEFSIYPAPQVSTA
VVEPYNSILTTHTTLEHSDCAFMVDNEAIYDICRRNLDIERPTYTNLNRLIGQIVSSITA
SLRFDGALNVDLTEFQTNLVPYPRAHFPLATYAPVISAEKAYHEQLSVAEITNACFEPAN
QMVKCDPRHGKYMACCLLYRGDVVPKDVNAAIATIKTKRTIQFVDWCPTGFKVGINYEPP
TVVPGGDLAKVQRAVCMLSNTTAIAEAWARLDHKFDLMYAKRAFVHWYVGEGMEEGEFSE
AREDMAALEKDYEEVGVDSVEGEGEEEGEEY
Ligands and cofactors
| ID | Name | Formula | Copies |
|---|
| GTP | Guanosine-5'-triphosphate | C10 H16 N5 O14 P3 | 2 |
| GDP | Guanosine-5'-diphosphate | C10 H15 N5 O11 P2 | 1 |
Primary citation
Cryo-EM structures of the tubulin cofactors reveal the molecular basis of alpha/beta-tubulin biogenesis. Taheri, A., Wang, Z., Singal, B. et al. Nat Commun (2025) 17:1405-1405. DOI 10.1038/s41467-025-68142-0 · PubMed
Other PDB entries of the same protein (UniProt P39937 (AlphaFold model), which also has an AlphaFold model), best resolution first:
- 9EDT 3.7 Å, Tubulin cofactors D,E,G bound to tubulin dimer
- 9EDR 3.8 Å, Tubulin Cofactors D,E,G,C and Tubulin complex -- TBCC N Terminus Bound to Tubulin
- 9EDS 3.8 Å, Tubulin cofactors D,E,G,C bound to tubulin dimer -- TBCC N terminus unbound
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