9M1N: Tubulin-specific chaperone C
Cryo-EM structure of the TBC-DC-Arl2-alpha-beta-tubulin complex with GDP-AlFx. Determined by electron microscopy at 2.24 Å resolution. Released 22 Oct 2025.
- Method
- Electron microscopy
- Resolution
- 2.24 Å
- Organisms
- Homo sapiens, Sus scrofa
- Chains
- 5
- Atoms
- 19,585
- Mol. weight
- 294.53 kDa
- Ligands
- GTP, MG, AF3, GDP
- Released
- 22 Oct 2025
Explore 9M1N in 3D
Show helices and sheets
RCSB PDB
PDBe
Secondary structure: helices and β-sheets
9M1N contains 133 α-helices and 67 β-strands across 5 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: 17 helices, 21 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 3-9 | 7 | 7 |
| α-helix | 11-28 | 18 | |
| α-helix | 49-51 | 3 | |
| β-strand | 53 | 1 | 8 |
| β-strand | 63 | 1 | 8 |
| β-strand | 67-69 | 3 | 7 |
| α-helix | 74-79 | 6 | |
| β-strand | 92-94 | 3 | 7 |
| α-helix | 103-107 | 5 | |
| α-helix | 115-127 | 13 | |
| β-strand | 132-133 | 2 | 7 |
| β-strand | 134 | 1 | 9 |
| β-strand | 135-138 | 4 | 7 |
| α-helix | 144-160 | 17 | |
| β-strand | 165 | 1 | 9 |
| β-strand | 168 | 1 | 7 |
| β-strand | 172 | 1 | 10 |
| α-helix | 183-194 | 12 | |
| β-strand | 201 | 1 | 7 |
| β-strand | 205 | 1 | 10 |
| α-helix | 206-215 | 10 | |
| α-helix | 224-238 | 15 | |
| α-helix | 252-259 | 8 | |
| β-strand | 269-273 | 5 | 11 |
| β-strand | 277 | 1 | 12 |
| α-helix | 288-293 | 6 | |
| α-helix | 294-296 | 3 | |
| β-strand | 301 | 1 | 11 |
| β-strand | 312-321 | 10 | 11 |
| α-helix | 325-338 | 14 | |
| β-strand | 343 | 1 | 11 |
| β-strand | 351-354 | 4 | 11 |
| β-strand | 368 | 1 | 12 |
| β-strand | 373-381 | 9 | 11 |
| α-helix | 382-384 | 3 | |
| α-helix | 385-399 | 15 | |
| α-helix | 406-409 | 4 | |
| α-helix | 416-435 | 20 | |
Chain b: 25 helices, 20 β-strands
| Element | Residues | Length | Sheet |
|---|
| β-strand | 3-8 | 6 | 13 |
| α-helix | 10-28 | 19 | |
| β-strand | 30 | 1 | 14 |
| β-strand | 36 | 1 | 14 |
| α-helix | 42-45 | 4 | |
| α-helix | 47-49 | 3 | |
| β-strand | 51-54 | 4 | 15 |
| α-helix | 55-57 | 3 | |
| β-strand | 58-61 | 4 | 15 |
| β-strand | 63-66 | 4 | 13 |
| α-helix | 71-77 | 7 | |
| α-helix | 82-84 | 3 | |
| α-helix | 87-89 | 3 | |
| β-strand | 90-91 | 2 | 13 |
| α-helix | 101-102 | 2 | |
| α-helix | 103-107 | 5 | |
| α-helix | 108-125 | 18 | |
| β-strand | 130-136 | 7 | 13 |
| β-strand | 138 | 1 | 16 |
| α-helix | 142-158 | 17 | |
| β-strand | 163-166 | 4 | 13 |
| β-strand | 169 | 1 | 16 |
| α-helix | 181-192 | 12 | |
| β-strand | 198-200 | 3 | 13 |
| 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 | 13 |
| β-strand | 267-271 | 5 | 17 |
| α-helix | 286-293 | 8 | |
| α-helix | 296-298 | 3 | |
| β-strand | 299 | 1 | 17 |
| β-strand | 310 | 1 | 18 |
| β-strand | 313-318 | 6 | 17 |
| α-helix | 323-336 | 14 | |
| β-strand | 341 | 1 | 18 |
| β-strand | 349-354 | 6 | 17 |
| α-helix | 357-358 | 2 | |
| β-strand | 364-369 | 6 | 17 |
| α-helix | 372-374 | 3 | |
| α-helix | 375-389 | 15 | |
| α-helix | 396-399 | 4 | |
| α-helix | 405-426 | 22 | |
Chain C: 11 helices, 12 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 22-24 | 3 | |
| α-helix | 27-50 | 24 | |
| α-helix | 53-55 | 3 | |
| α-helix | 58-76 | 19 | |
| α-helix | 81-99 | 19 | |
| α-helix | 107-131 | 25 | |
| α-helix | 134-137 | 4 | |
| β-strand | 187-190 | 4 | 1 |
| β-strand | 196 | 1 | 1 |
| α-helix | 198-200 | 3 | |
| β-strand | 206-212 | 7 | 1 |
| β-strand | 215-218 | 4 | 1 |
| β-strand | 225-231 | 7 | 1 |
| β-strand | 234-237 | 4 | 1 |
| β-strand | 243-255 | 13 | 1 |
| β-strand | 257-272 | 16 | 1 |
| β-strand | 274-285 | 12 | 1 |
| β-strand | 287-290 | 4 | 1 |
| α-helix | 299-303 | 5 | |
| α-helix | 304-306 | 3 | |
| β-strand | 319-320 | 2 | 1 |
| β-strand | 333-335 | 3 | 1 |
| α-helix | 338-340 | 3 | |
Chain D: 73 helices, 6 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 33-40 | 8 | |
| α-helix | 54-65 | 12 | |
| α-helix | 71-74 | 4 | |
| α-helix | 80-89 | 10 | |
| α-helix | 96-112 | 17 | |
| α-helix | 115-118 | 4 | |
| α-helix | 119-121 | 3 | |
| α-helix | 130-137 | 8 | |
| α-helix | 144-146 | 3 | |
| α-helix | 147-159 | 13 | |
| α-helix | 167-171 | 5 | |
| α-helix | 181-184 | 4 | |
| α-helix | 185-196 | 12 | |
| α-helix | 203-214 | 12 | |
| α-helix | 218-224 | 7 | |
| α-helix | 226-237 | 12 | |
| α-helix | 244-263 | 20 | |
| α-helix | 266-282 | 17 | |
| α-helix | 285-287 | 3 | |
| α-helix | 291-306 | 16 | |
| α-helix | 357-369 | 13 | |
| α-helix | 374-388 | 15 | |
| α-helix | 395-404 | 10 | |
| α-helix | 405-407 | 3 | |
| α-helix | 413-428 | 16 | |
| α-helix | 434-436 | 3 | |
| α-helix | 437-447 | 11 | |
| β-strand | 452-453 | 2 | 2 |
| β-strand | 458-459 | 2 | 2 |
| α-helix | 461-477 | 17 | |
| α-helix | 484-486 | 3 | |
| α-helix | 487-500 | 14 | |
| α-helix | 504-520 | 17 | |
| α-helix | 527-533 | 7 | |
| α-helix | 538-540 | 3 | |
| α-helix | 542-554 | 13 | |
| α-helix | 561-569 | 9 | |
| α-helix | 571-573 | 3 | |
| α-helix | 577-589 | 13 | |
| α-helix | 595-597 | 3 | |
| α-helix | 598-602 | 5 | |
| α-helix | 603-609 | 7 | |
| α-helix | 615-637 | 23 | |
| α-helix | 649-655 | 7 | |
| α-helix | 658-664 | 7 | |
| α-helix | 671-689 | 19 | |
| α-helix | 698-712 | 15 | |
| α-helix | 714-716 | 3 | |
| α-helix | 719-721 | 3 | |
| α-helix | 722-736 | 15 | |
| α-helix | 750-761 | 12 | |
| α-helix | 769-779 | 11 | |
| α-helix | 791-801 | 11 | |
| α-helix | 812-829 | 18 | |
| β-strand | 831 | 1 | 3 |
| β-strand | 840 | 1 | 3 |
| α-helix | 845-854 | 10 | |
| β-strand | 861-862 | 2 | 4 |
| β-strand | 865-866 | 2 | 4 |
| α-helix | 868-887 | 20 | |
| α-helix | 891-893 | 3 | |
| α-helix | 896-909 | 14 | |
| α-helix | 915-929 | 15 | |
| α-helix | 935-936 | 2 | |
| α-helix | 942-948 | 7 | |
| α-helix | 966-969 | 4 | |
| α-helix | 970-974 | 5 | |
| α-helix | 976-990 | 15 | |
| α-helix | 995-1013 | 19 | |
| α-helix | 1016-1019 | 4 | |
| α-helix | 1022-1032 | 11 | |
| α-helix | 1040-1053 | 14 | |
| α-helix | 1066-1079 | 14 | |
| α-helix | 1085-1098 | 14 | |
| α-helix | 1104-1117 | 14 | |
| α-helix | 1122-1138 | 17 | |
| α-helix | 1148-1157 | 10 | |
| α-helix | 1164-1178 | 15 | |
| α-helix | 1181-1184 | 4 | |
Chain G: 7 helices, 8 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 3-14 | 12 | |
| β-strand | 16-23 | 8 | 5 |
| α-helix | 29-37 | 9 | |
| β-strand | 50-57 | 8 | 5 |
| β-strand | 60-67 | 8 | 5 |
| α-helix | 71-74 | 4 | |
| α-helix | 77-80 | 4 | |
| β-strand | 86-92 | 7 | 5 |
| α-helix | 99-108 | 10 | |
| β-strand | 119-125 | 7 | 5 |
| α-helix | 135-141 | 7 | |
| β-strand | 154-156 | 3 | 5 |
| β-strand | 158 | 1 | 6 |
| β-strand | 163 | 1 | 6 |
| α-helix | 166-178 | 13 | |
Molecules and chains
| Molecule | Chains | Type | Length | Organism | UniProt |
|---|
| Tubulin-specific chaperone C | C | protein | 346 | Homo sapiens | Q15814 (AlphaFold model) |
| Tubulin-specific chaperone D | D | protein | 1192 | Homo sapiens | Q9BTW9 (AlphaFold model) |
| ADP-ribosylation factor-like protein 2 | G | protein | 184 | Homo sapiens | P36404 (AlphaFold model) |
| Tubulin alpha-1B chain | a | protein | 451 | Sus scrofa | Q2XVP4 (AlphaFold model) |
| Tubulin beta chain | b | protein | 444 | Sus scrofa | Q767L7 |
Sequence of entity 1 (C), FASTA
>9M1N_1 Tubulin-specific chaperone C (chains C)
MESVSCSAAAVRTGDMESQRDLSLVPERLQRREQERQLEVERRKQKRQNQEVEKENSHFF
VATFVRERAAVEELLERAESVERLEEAASRLQGLQKLINDSVFFLAAYDLRQGQEALARL
QAALAERRRGLQPKKRFAFKTRGKDAASSTKVDAAPGIPPAVESIQDSPLPKKAEGDLGP
SWVCGFSNLESQVLEKRASELHQRDVLLTELSNCTVRLYGNPNTLRLTKAHSCKLLCGPV
STSVFLEDCSDCVLAVACQQLRIHSTKDTRIFLQVTSRAIVEDCSGIQFAPYTWSYPEID
KDFESSGLDRSKNNWNDVDDFNWLARDMASPNWSILPEEERNIQWD
Sequence of entity 2 (D), FASTA
>9M1N_2 Tubulin-specific chaperone D (chains D)
MALSDEPAAGGPEEEAEDETLAFGAALEAFGESAETRALLGRLREVHGGGAEREVALERF
RVIMDKYQEQPHLLDPHLEWMMNLLLDIVQDQTSPASLVHLAFKFLYIITKVRGYKTFLR
LFPHEVADVEPVLDLVTIQNPKDHEAWETRYMLLLWLSVTCLIPFDFSRLDGNLLTQPGQ
ARMSIMDRILQIAESYLIVSDKARDAAAVLVSRFITRPDVKQSKMAEFLDWSLCNLARSS
FQTMQGVITMDGTLQALAQIFKHGKREDCLPYAATVLRCLDGCRLPESNQTLLRKLGVKL
VQRLGLTFLKPKVAAWRYQRGCRSLAANLQLLTQGQSEQKPLILTEDDDEDDDVPEGVER
VIEQLLVGLKDKDTVVRWSAAKGIGRMAGRLPRALADDVVGSVLDCFSFQETDKAWHGGC
LALAELGRRGLLLPSRLVDVVAVILKALTYDEKRGACSVGTNVRDAACYVCWAFARAYEP
QELKPFVTAISSALVIAAVFDRDINCRRAASAAFQENVGRQGTFPHGIDILTTADYFAVG
NRSNCFLVISVFIAGFPEYTQPMIDHLVTMKISHWDGVIRELAARALHNLAQQAPEFSAT
QVFPRLLSMTLSPDLHMRHGSILACAEVAYALYKLAAQENRPVTDHLDEQAVQGLKQIHQ
QLYDRQLYRGLGGQLMRQAVCVLIEKLSLSKMPFRGDTVIDGWQWLINDTLRHLHLISSH
SRQQMKDAAVSALAALCSEYYMKEPGEADPAIQEELITQYLAELRNPEEMTRCGFSLALG
ALPGFLLKGRLQQVLTGLRAVTHTSPEDVSFAESRRDGLKAIARICQTVGVKAGAPDEAV
CGENVSQIYCALLGCMDDYTTDSRGDVGTWVRKAAMTSLMDLTLLLARSQPELIEAHTCE
RIMCCVAQQASEKIDRFRAHAASVFLTLLHFDSPPIPHVPHRGELEKLFPRSDVASVNWS
APSQAFPRITQLLGLPTYRYHVLLGLVVSLGGLTESTIRHSTQSLFEYMKGIQSDPQALG
SFSGTLLQIFEDNLLNERVSVPLLKTLDHVLTHGCFDIFTTEEDHPFAVKLLALCKKEIK
NSKDIQKLLSGIAVFCEMVQFPGDVRRQALLQLCLLLCHRFPLIRKTTASQVYETLLTYS
DVVGADVLDEVVTVLSDTAWDAELAVVREQRNRLCDLLGVPRPQLVPQPGAC
Sequence of entity 3 (G), FASTA
>9M1N_3 ADP-ribosylation factor-like protein 2 (chains G)
MGLLTILKKMKQKERELRLLMLGLDNAGKTTILKKFNGEDIDTISPTLGFNIKTLEHRGF
KLNIWDVGGQKSLRSYWRNYFESTDGLIWVVDSADRQRMQDCQRELQSLLVEERLAGATL
LIFANKQDLPGALSSNAIREVLELDSIRSHHWCIQGCSAVTGENLLPGIDWLLDDISSRI
FTAD
Sequence of entity 4 (a), FASTA
>9M1N_4 Tubulin alpha-1B chain (chains a)
MRECISIHVGQAGVQIGNACWELYCLEHGIQPDGQMPSDKTIGGGDDSFNTFFSETGAGK
HVPRAVFVDLEPTVIDEVRTGTYRQLFHPEQLITGKEDAANNYARGHYTIGKEIIDLVLD
RIRKLADQCTGLQGFLVFHSFGGGTGSGFTSLLMERLSVDYGKKSKLEFSIYPAPQVSTA
VVEPYNSILTTHTTLEHSDCAFMVDNEAIYDICRRNLDIERPTYTNLNRLISQIVSSITA
SLRFDGALNVDLTEFQTNLVPYPRIHFPLATYAPVISAEKAYHEQLSVAEITNACFEPAN
QMVKCDPRHGKYMACCLLYRGDVVPKDVNAAIATIKTKRSIQFVDWCPTGFKVGINYQPP
TVVPGGDLAKVQRAVCMLSNTTAIAEAWARLDHKFDLMYAKRAFVHWYVGEGMEEGEFSE
AREDMAALEKDYEEVGVDSVEGEGEEEGEEY
Sequence of entity 5 (b), FASTA
>9M1N_5 Tubulin beta chain (chains b)
MREIVHIQAGQCGNQIGAKFWEVISDEHGIDPTGTYHGDSDLQLDRISVYYNEATGGKYV
PRAILVDLEPGTMDSVRSGPFGQIFRPDNFVFGQSGAGNNWAKGHYTEGAELVDSVLDVV
RKEAESCDCLQGFQLTHSLGGGTGSGMGTLLISKIREEYPDRIMNTFSVVPSPKVSDTVV
EPYNATLSVHQLVENTDETYCIDNEALYDICFRTLKLTTPTYGDLNHLVSATMSGVTTCL
RFPGQLNADLRKLAVNMVPFPRLHFFMPGFAPLTSRGSQQYRALTVPELTQQVFDAKNMM
AACDPRHGRYLTVAAVFRGRMSMKEVDEQMLNVQNKNSSYFVEWIPNNVKTAVCDIPPRG
LKMAVTFIGNSTAIQELFKRISEQFTAMFRRKAFLHWYTGEGMDEMEFTEAESNMNDLVS
EYQQYQDATAEEEEDFGEEAEEEA
Ligands and cofactors
| ID | Name | Formula | Copies |
|---|
| GTP | Guanosine-5'-triphosphate | C10 H16 N5 O14 P3 | 2 |
| MG | Magnesium ion | Mg | 3 |
| AF3 | Aluminum fluoride | Al F3 | 1 |
| GDP | Guanosine-5'-diphosphate | C10 H15 N5 O11 P2 | 1 |
Primary citation
Structural dissection of alpha beta-tubulin heterodimer assembly and disassembly by human tubulin-specific chaperones. Seong, Y., Kim, H., Byun, K. et al. Science (2025) 390:eady2708-eady2708. DOI 10.1126/science.ady2708 · PubMed
Other PDB entries of the same protein (UniProt Q15814 (AlphaFold model), which also has an AlphaFold model), best resolution first:
- 9M1M 2.21 Å, Cryo-EM structure of the TBC-DEC-Arl2-alpha-beta-tubulin complex with GDP-AlFx
- 2L3L The solution structure of the N-terminal domain of human Tubulin Binding Cofactor C…
- 2YUH Solution structure of the C-terminal region in human tubulin folding cofactor C
Browse structure collections
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