9EDS: Tubulin cofactors D,E,G,C
Tubulin cofactors D,E,G,C bound to tubulin dimer -- TBCC N terminus unbound. Determined by electron microscopy at 3.8 Å resolution. Released 4 Feb 2026.
- Method
- Electron microscopy
- Resolution
- 3.8 Å
- Organisms
- Saccharomyces cerevisiae, Sus scrofa, Escherichia coli
- Chains
- 7
- Atoms
- 20,957
- Mol. weight
- 333.72 kDa
- Ligands
- GDP, GTP
- Released
- 4 Feb 2026
Explore 9EDS in 3D
Show helices and sheets
RCSB PDB
PDBe
Secondary structure: helices and β-sheets
9EDS contains 160 α-helices and 96 β-strands across 7 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: 27 helices, 14 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 3-9 | 7 | 17 |
| α-helix | 10-28 | 19 | |
| α-helix | 49-51 | 3 | |
| β-strand | 53-55 | 3 | 18 |
| β-strand | 61-63 | 3 | 18 |
| β-strand | 65-69 | 5 | 17 |
| α-helix | 72-80 | 9 | |
| α-helix | 82-85 | 4 | |
| α-helix | 89-91 | 3 | |
| β-strand | 92-94 | 3 | 17 |
| α-helix | 103-108 | 6 | |
| α-helix | 110-127 | 18 | |
| β-strand | 132-140 | 9 | 17 |
| α-helix | 144 | 1 | |
| α-helix | 145-149 | 5 | |
| α-helix | 150-161 | 12 | |
| β-strand | 165-172 | 8 | 17 |
| α-helix | 173 | 1 | |
| α-helix | 175-177 | 3 | |
| α-helix | 183-195 | 13 | |
| β-strand | 200-205 | 6 | 17 |
| α-helix | 206-216 | 11 | |
| α-helix | 224-243 | 20 | |
| α-helix | 252-259 | 8 | |
| α-helix | 268 | 1 | |
| β-strand | 269-273 | 5 | 19 |
| α-helix | 280-282 | 3 | |
| α-helix | 288-295 | 8 | |
| β-strand | 301 | 1 | 19 |
| α-helix | 307-309 | 3 | |
| β-strand | 312-321 | 10 | 19 |
| α-helix | 325-338 | 14 | |
| α-helix | 342 | 1 | |
| β-strand | 343 | 1 | 19 |
| α-helix | 344 | 1 | |
| β-strand | 352-356 | 5 | 19 |
| α-helix | 359-360 | 2 | |
| β-strand | 373-381 | 9 | 19 |
| α-helix | 384-400 | 17 | |
| α-helix | 405-410 | 6 | |
| α-helix | 416-435 | 20 | |
Chain B: 25 helices, 19 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 3-8 | 6 | 20 |
| α-helix | 10-28 | 19 | |
| β-strand | 30 | 1 | 21 |
| β-strand | 35 | 1 | 22 |
| β-strand | 36 | 1 | 21 |
| α-helix | 42-45 | 4 | |
| α-helix | 47-49 | 3 | |
| β-strand | 51-54 | 4 | 22 |
| β-strand | 58-61 | 4 | 22 |
| β-strand | 63-65 | 3 | 20 |
| β-strand | 66-67 | 2 | 23 |
| α-helix | 70-78 | 9 | |
| α-helix | 87-89 | 3 | |
| β-strand | 91-92 | 2 | 23 |
| α-helix | 101-106 | 6 | |
| α-helix | 108-124 | 17 | |
| β-strand | 130-138 | 9 | 20 |
| α-helix | 143-158 | 16 | |
| β-strand | 163-170 | 8 | 20 |
| α-helix | 171 | 1 | |
| α-helix | 173-175 | 3 | |
| α-helix | 181-195 | 15 | |
| β-strand | 198-203 | 6 | 20 |
| α-helix | 204-209 | 6 | |
| α-helix | 210-214 | 5 | |
| α-helix | 222-236 | 15 | |
| α-helix | 238-241 | 4 | |
| α-helix | 250-257 | 8 | |
| β-strand | 265-266 | 2 | 20 |
| β-strand | 267-271 | 5 | 24 |
| α-helix | 286-294 | 9 | |
| α-helix | 296-298 | 3 | |
| β-strand | 299 | 1 | 24 |
| α-helix | 305-307 | 3 | |
| β-strand | 310-319 | 10 | 24 |
| α-helix | 323-336 | 14 | |
| α-helix | 338-340 | 3 | |
| β-strand | 341 | 1 | 24 |
| β-strand | 349-354 | 6 | 24 |
| α-helix | 357-358 | 2 | |
| β-strand | 363-371 | 9 | 24 |
| α-helix | 374-389 | 16 | |
| α-helix | 395-399 | 5 | |
| α-helix | 405-424 | 20 | |
Chain C: 5 helices, 16 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 103-104 | 2 | 25 |
| β-strand | 107-110 | 4 | 26 |
| β-strand | 116-121 | 6 | 25 |
| β-strand | 124-127 | 4 | 26 |
| α-helix | 134-135 | 2 | |
| β-strand | 136-147 | 12 | 25 |
| β-strand | 150-153 | 4 | 26 |
| β-strand | 158-169 | 12 | 25 |
| β-strand | 172-177 | 6 | 26 |
| β-strand | 183-188 | 6 | 25 |
| β-strand | 191 | 1 | 25 |
| β-strand | 194-199 | 6 | 26 |
| α-helix | 207 | 1 | |
| β-strand | 208-213 | 6 | 25 |
| β-strand | 216 | 1 | 25 |
| β-strand | 218-221 | 4 | 26 |
| α-helix | 222-224 | 3 | |
| β-strand | 228-232 | 5 | 25 |
| β-strand | 248-251 | 4 | 26 |
| α-helix | 256-258 | 3 | |
| α-helix | 260-267 | 8 | |
Chain D: 66 helices, 6 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 2-16 | 15 | |
| α-helix | 20-31 | 12 | |
| α-helix | 32-35 | 4 | |
| α-helix | 37-40 | 4 | |
| α-helix | 44-53 | 10 | |
| α-helix | 55-57 | 3 | |
| α-helix | 60-83 | 24 | |
| α-helix | 91-101 | 11 | |
| α-helix | 108-125 | 18 | |
| α-helix | 132-142 | 11 | |
| α-helix | 152-166 | 15 | |
| α-helix | 171-173 | 3 | |
| α-helix | 176-179 | 4 | |
| α-helix | 182-192 | 11 | |
| α-helix | 196-202 | 7 | |
| α-helix | 205-217 | 13 | |
| α-helix | 224-234 | 11 | |
| α-helix | 237-249 | 13 | |
| α-helix | 256-276 | 21 | |
| α-helix | 278-295 | 18 | |
| α-helix | 305-320 | 16 | |
| α-helix | 326-335 | 10 | |
| α-helix | 337-340 | 4 | |
| β-strand | 345-347 | 3 | 1 |
| β-strand | 350-352 | 3 | 1 |
| α-helix | 354-369 | 16 | |
| α-helix | 378-394 | 17 | |
| α-helix | 398-414 | 17 | |
| α-helix | 416-419 | 4 | |
| α-helix | 423-431 | 9 | |
| α-helix | 437-451 | 15 | |
| 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 | 505-513 | 9 | |
| α-helix | 519-534 | 16 | |
| α-helix | 542-553 | 12 | |
| α-helix | 571-586 | 16 | |
| α-helix | 593-603 | 11 | |
| α-helix | 612-625 | 14 | |
| α-helix | 631-639 | 9 | |
| α-helix | 643-648 | 6 | |
| α-helix | 649-651 | 3 | |
| α-helix | 655-665 | 11 | |
| α-helix | 671-684 | 14 | |
| α-helix | 685-689 | 5 | |
| α-helix | 694-707 | 14 | |
| β-strand | 714-715 | 2 | 2 |
| β-strand | 717 | 1 | 2 |
| 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 | 830-843 | 14 | |
| α-helix | 845-846 | 2 | |
| α-helix | 847-853 | 7 | |
| α-helix | 856-859 | 4 | |
| β-strand | 860 | 1 | 3 |
| α-helix | 866 | 1 | |
| β-strand | 867 | 1 | 3 |
| 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 | 989-993 | 5 | |
Chain E: 15 helices, 33 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 7-11 | 5 | 4 |
| β-strand | 14-18 | 5 | 4 |
| β-strand | 20-23 | 4 | 5 |
| β-strand | 31-34 | 4 | 5 |
| β-strand | 45 | 1 | 6 |
| β-strand | 48-49 | 2 | 7 |
| β-strand | 52-53 | 2 | 7 |
| β-strand | 63 | 1 | 6 |
| β-strand | 67 | 1 | 5 |
| α-helix | 68-71 | 4 | |
| β-strand | 126-128 | 3 | 8 |
| α-helix | 143-147 | 5 | |
| α-helix | 148-153 | 6 | |
| β-strand | 156-158 | 3 | 8 |
| β-strand | 166 | 1 | 9 |
| α-helix | 168-174 | 7 | |
| β-strand | 182-184 | 3 | 8 |
| β-strand | 192 | 1 | 9 |
| β-strand | 207-209 | 3 | 8 |
| α-helix | 217-224 | 8 | |
| β-strand | 232-234 | 3 | 8 |
| β-strand | 241 | 1 | 10 |
| α-helix | 244-248 | 5 | |
| β-strand | 258-260 | 3 | 8 |
| β-strand | 265 | 1 | 10 |
| β-strand | 280-282 | 3 | 8 |
| β-strand | 301-303 | 3 | 8 |
| β-strand | 311 | 1 | 11 |
| α-helix | 314-321 | 8 | |
| β-strand | 336-337 | 2 | 11 |
| β-strand | 340-341 | 2 | 11 |
| α-helix | 347-356 | 10 | |
| β-strand | 362-363 | 2 | 12 |
| β-strand | 366-367 | 2 | 12 |
| α-helix | 370-383 | 14 | |
| α-helix | 384-386 | 3 | |
| α-helix | 396-405 | 10 | |
| α-helix | 409-412 | 4 | |
| β-strand | 426-431 | 6 | 13 |
| α-helix | 436-438 | 3 | |
| β-strand | 439-444 | 6 | 13 |
| β-strand | 449 | 1 | 14 |
| α-helix | 450-461 | 12 | |
| β-strand | 469-473 | 5 | 13 |
| β-strand | 479-481 | 3 | 13 |
| β-strand | 488 | 1 | 14 |
| α-helix | 489-492 | 4 | |
| β-strand | 499-503 | 5 | 13 |
Chain G: 9 helices, 8 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 3-14 | 12 | |
| β-strand | 16-22 | 7 | 15 |
| α-helix | 29-36 | 8 | |
| α-helix | 39-41 | 3 | |
| β-strand | 53-60 | 8 | 15 |
| β-strand | 63-70 | 8 | 15 |
| α-helix | 74-76 | 3 | |
| α-helix | 78-83 | 6 | |
| β-strand | 89-95 | 7 | 15 |
| α-helix | 99-101 | 3 | |
| α-helix | 102-115 | 14 | |
| β-strand | 120 | 1 | 16 |
| β-strand | 124 | 1 | 16 |
| β-strand | 125-131 | 7 | 15 |
| α-helix | 139-156 | 18 | |
| β-strand | 164-167 | 4 | 15 |
| α-helix | 176-190 | 15 | |
Chain I: 13 helices, 0 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 15-22 | 8 | |
| α-helix | 28-41 | 14 | |
| α-helix | 44-46 | 3 | |
| α-helix | 47-53 | 7 | |
| α-helix | 59-72 | 14 | |
| α-helix | 75-77 | 3 | |
| α-helix | 78-84 | 7 | |
| α-helix | 90-103 | 14 | |
| α-helix | 106-108 | 3 | |
| α-helix | 109-115 | 7 | |
| α-helix | 121-134 | 14 | |
| α-helix | 138-148 | 11 | |
| α-helix | 152-164 | 13 | |
Molecules and chains
| Molecule | Chains | Type | Length | Organism | UniProt |
|---|
| Chromosome instability protein 1 | D | protein | 1021 | Saccharomyces cerevisiae | P40987 (AlphaFold model) |
| Protein PAC2 | E | protein | 469 | Saccharomyces cerevisiae | P39937 (AlphaFold model) |
| GTP-binding protein CIN4 | G | protein | 191 | Saccharomyces cerevisiae | P39110 (AlphaFold model) |
| Tubulin alpha-1A chain | A | protein | 451 | Sus scrofa | P02550 (AlphaFold model) |
| Tubulin beta chain | B | protein | 445 | Sus scrofa | P02554 |
| iih5 Alpha Repeating | I | protein | 170 | Escherichia coli | |
| Tubulin-specific chaperone C | C | protein | 268 | Saccharomyces cerevisiae | P46670 |
Sequence of entity 1 (D), FASTA
>9EDS_1 Chromosome instability protein 1 (chains D)
MHHHHHHMNNIRALLDSIQSGVQTVSPEKHQQTIAAINKFQDDPALLDTILPRCVPLLTK
SFFCMSQRDQKLVAELFYNLDKISHSKVLKSLDTSIFRLNEILNYLQDRASPSSFSDVLC
VYLNLSWLSVILLSPYAFKDKFNKTLQVSSRFENYPICIPPINKIKAVLYFKNFTDAFDQ
LPEREQANVPFLNQFLKLFIQSSEKANYYFSNENLRHLQQVALSNDGIKLLPKLFQISFN
HGSHDILDAIVEFFHDHLSSNSTDTRFQLAHSFAKIAKFLHQADPASFIELIDYTIENTV
SLLQAPCDSIDSNELHTSLLIIAEVALAKILPIDLVDRVLTLIIPKTCHFQKSHFQIIKG
HHIRDSTNFIIWSVIRSNRSNSLSPQVLQSLLSHLLINAFFDPELIIRYSSFAALQELLG
RSNKSLALNQNDIALILQANWKDLPRSFEENSGLIRRLFNPENTSKSAVCVWKVFRDWSF
NWNLLENLHLTTMKLNIDYNLVPLIKSKLSSPALLQEVLNKAGSSVTQNCQILYLYLKLF
ENDVNCPKISEICIDIYQKKIKFQLTTQAKRQFNDNSPELFQIFVILKYWQLTGQNDFNQ
ELFWKFVDIVSPQKKLNLYNEFIPIIQQIISQCVSLNYTRIVQLIKSDNELTCRSICHMP
DQEKMCSLFFSQFPLLSPQSRSLLIGELDHHWDVRISLLPSNSYRKFRNIIINCLDDYTI
TQQGDVGRLVRIQALKLMQSHPDFLSGDCDSINPKLTRLLAEPVPEIRKLSYQLLASATS
QITVLSDSSILNFRHKQGLSEEFWKGYAVSAGAIHFTDSQLTSSIDSFIVYYRSLSPSQQ
LELCHDLIRIIPSAKQIAESRIRDRNKDPLTGGMRFDTIKFTIHCVKFWTRIMESGLVVL
HPNFNFQGVFAKFYNLHLLDCTTLRVSVIKFFPFLAISCYHTMRENADQKNLSNIILKRL
LVLVKREYAATKSKFMTDQNVALQGMFQIFLELGVTRQLQALQVACQKHELANILESDIT
L
Sequence of entity 2 (E), FASTA
>9EDS_2 Protein PAC2 (chains E)
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
XXXXXXXXXXXXQSLVLKIKIRAGKKPSSDLDYWVLPSFTVRYVKSVICRKLNFDILNVK
LFHENSEGMINEIKYNFRPISDFNVVNGDIIHVSSPVNNKSIQKVNSPS
Sequence of entity 3 (G), FASTA
>9EDS_3 GTP-binding protein CIN4 (chains G)
MGLLSIIRKQKLRDKEIRCLILGLDNSGKSTIVNKLLPKDEQNNDGIMPTVGFQIHSLMI
KDVTISLWDIGGLRTLRPFWDNYFDKTQAMIWCIDVSLSMRFDETLQELKELINRDENRI
GYECAVIVVLNKIDLVEDKSELHRRCLLVESELKCLFKPDIRIELVKCSGVTGEGIDNLR
DRLVESCHFTQ
Sequence of entity 4 (A), FASTA
>9EDS_4 Tubulin alpha-1A chain (chains A)
MRECISIHVGQAGVQIGNACWELYCLEHGIQPDGQMPSDKTIGGGDDSFNTFFSETGAGK
HVPRAVFVDLEPTVIDEVRTGTYRQLFHPEQLITGKEDAANNYARGHYTIGKEIIDLVLD
RIRKLADQCTGLQGFSVFHSFGGGTGSGFTSLLMERLSVDYGKKSKLEFSIYPAPQVSTA
VVEPYNSILTTHTTLEHSDCAFMVDNEAIYDICRRNLDIERPTYTNLNRLIGQIVSSITA
SLRFDGALNVDLTEFQTNLVPYPRAHFPLATYAPVISAEKAYHEQLSVAEITNACFEPAN
QMVKCDPRHGKYMACCLLYRGDVVPKDVNAAIATIKTKRTIQFVDWCPTGFKVGINYEPP
TVVPGGDLAKVQRAVCMLSNTTAIAEAWARLDHKFDLMYAKRAFVHWYVGEGMEEGEFSE
AREDMAALEKDYEEVGVDSVEGEGEEEGEEY
Sequence of entity 5 (B), FASTA
>9EDS_5 Tubulin beta chain (chains B)
MREIVHIQAGQCGNQIGAKFWEVISDEHGIDPTGSYHGDSDLQLERINVYYNEAAGNKYV
PRAILVDLEPGTMDSVRSGPFGQIFRPDNFVFGQSGAGNNWAKGHYTEGAELVDSVLDVV
RKESESCDCLQGFQLTHSLGGGTGSGMGTLLISKIREEYPDRIMNTFSVVPSPKVSDTVV
EPYNATLSVHQLVENTDETYCIDNEALYDICFRTLKLTTPTYGDLNHLVSATMSGVTTCL
RFPGQLNADLRKLAVNMVPFPRLHFFMPGFAPLTSRGSQQYRALTVPELTQQMFDAKNMM
AACDPRHGRYLTVAAVFRGRMSMKEVDEQMLNVQNKNSSYFVEWIPNNVKTAVCDIPPRG
LKMSATFIGNSTAIQELFKRISEQFTAMFRRKAFLHWYTGEGMDEMEFTEAESNMNDLVS
EYQQYQDATADEQGEFEEEGEEDEA
Sequence of entity 6 (I), FASTA
>9EDS_6 iih5 Alpha Repeating (chains I)
MRGSHHHHHHTDPEKVEMYIKNLQDDSPPVRFNAAVALGKIGDERAVEPLIKALKDEDWQ
VRKTAAYALGKIGDERAVEPLIKALKDEDRYVRSRAALALGKIGDERAVEPLIKALKDED
EYVRLSAASALGKIGGERVRAAMEKLAETGTGFARKVAVNYLETHKSLIS
Sequence of entity 7 (C), FASTA
>9EDS_7 Tubulin-specific chaperone C (chains C)
MDFTAKIKELERELSETSDYKTLQKKTISLRSELNTLSHSLTSYEKEHFSNDIENVLKSI
NAKLSESKGKKRLFSFKQKNSSSAVHKNVERTELANAPAYTTTLKKHYVLEKGDSAFENL
EFCTVTSTTDYSGNSALSGSLCFRNITKCVINLQRIFFQTGSIFITDCTDSIIFLRSPSD
KDFQIRLRDLKNCKILIEKLSPSIDCKQVVIIENCHKCIFNASTRDHLIIQDFSNPFQSE
ETEDNSAFAFEDFDICNKDTMQLFRAYL
Ligands and cofactors
| ID | Name | Formula | Copies |
|---|
| GDP | Guanosine-5'-diphosphate | C10 H15 N5 O11 P2 | 1 |
| GTP | Guanosine-5'-triphosphate | C10 H16 N5 O14 P3 | 2 |
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 P40987 (AlphaFold model), which also has an AlphaFold model), best resolution first:
- 9EDT 3.7 Å, Tubulin cofactors D,E,G bound to tubulin dimer
- 9EEB 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
Browse structure collections
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