9P73: In situ human eEF2-A/P-P/E state 80S ribosome
In situ human eEF2-A/P-P/E state 80S ribosome. Determined by electron microscopy at 2.66 Å resolution. Released 24 Dec 2025.
- Method
- Electron microscopy
- Resolution
- 2.66 Å
- Organism
- Homo sapiens
- Chains
- 85
- Atoms
- 227,131
- Mol. weight
- 3359.9 kDa
- Ligands
- MG, ZN, SPM, SPD
- Released
- 24 Dec 2025
Explore 9P73 in 3D
Show helices and sheets
RCSB PDB
PDBe
Secondary structure: helices and β-sheets
9P73 contains 595 α-helices and 612 β-strands across 79 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 CB: 37 helices, 46 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 7-13 | 7 | |
| β-strand | 19-26 | 8 | 10 |
| α-helix | 32-42 | 11 | |
| β-strand | 73-80 | 8 | 10 |
| α-helix | 81-82 | 2 | |
| α-helix | 83-88 | 6 | |
| β-strand | 97-104 | 8 | 10 |
| α-helix | 114-121 | 8 | |
| β-strand | 124-130 | 7 | 10 |
| β-strand | 134 | 1 | 10 |
| α-helix | 137-148 | 12 | |
| β-strand | 152-158 | 7 | 10 |
| α-helix | 160-161 | 2 | |
| α-helix | 162-166 | 5 | |
| α-helix | 171-192 | 22 | |
| β-strand | 212-216 | 5 | 10 |
| β-strand | 221-223 | 3 | 10 |
| α-helix | 225-235 | 11 | |
| α-helix | 248-259 | 12 | |
| β-strand | 265-267 | 3 | 11 |
| β-strand | 272-274 | 3 | 11 |
| β-strand | 278 | 1 | 12 |
| β-strand | 284 | 1 | 12 |
| β-strand | 287 | 1 | 11 |
| α-helix | 288-289 | 2 | |
| α-helix | 290-294 | 5 | |
| α-helix | 295-305 | 11 | |
| α-helix | 309-319 | 11 | |
| α-helix | 333-343 | 11 | |
| β-strand | 345 | 1 | 10 |
| α-helix | 346-357 | 12 | |
| α-helix | 359-360 | 2 | |
| α-helix | 361-372 | 12 | |
| β-strand | 373 | 1 | 13 |
| α-helix | 380-387 | 8 | |
| β-strand | 395-403 | 9 | 14 |
| β-strand | 410-418 | 9 | 14 |
| β-strand | 420-421 | 2 | 15 |
| β-strand | 426-430 | 5 | 14 |
| β-strand | 442-446 | 5 | 14 |
| β-strand | 449-454 | 6 | 14 |
| β-strand | 457-461 | 5 | 14 |
| β-strand | 464-465 | 2 | 15 |
| β-strand | 468-473 | 6 | 14 |
| α-helix | 476-478 | 3 | |
| β-strand | 480 | 1 | 16 |
| β-strand | 482 | 1 | 16 |
| β-strand | 483-486 | 4 | 14 |
| β-strand | 494 | 1 | 13 |
| α-helix | 495-497 | 3 | |
| β-strand | 505-511 | 7 | 17 |
| α-helix | 517-530 | 14 | |
| β-strand | 535-538 | 4 | 17 |
| β-strand | 544-548 | 5 | 17 |
| α-helix | 551-560 | 10 | |
| α-helix | 561-565 | 5 | |
| β-strand | 570-573 | 4 | 17 |
| α-helix | 574-578 | 5 | |
| β-strand | 581-583 | 3 | 18 |
| β-strand | 591-594 | 4 | 19 |
| β-strand | 601-608 | 8 | 19 |
| α-helix | 613-619 | 7 | |
| α-helix | 628-638 | 11 | |
| α-helix | 643-647 | 5 | |
| β-strand | 649-651 | 3 | 19 |
| α-helix | 654-656 | 3 | |
| β-strand | 660-664 | 5 | 19 |
| α-helix | 672-688 | 17 | |
| β-strand | 697 | 1 | 18 |
| β-strand | 700-708 | 9 | 19 |
| α-helix | 718-720 | 3 | |
| α-helix | 722-735 | 14 | |
| β-strand | 738-740 | 3 | 18 |
| β-strand | 741-742 | 2 | 20 |
| β-strand | 744-751 | 8 | 21 |
| α-helix | 756-765 | 10 | |
| β-strand | 769-775 | 7 | 21 |
| β-strand | 782-789 | 8 | 21 |
| α-helix | 796-804 | 9 | |
| β-strand | 809-814 | 6 | 21 |
| β-strand | 818-819 | 2 | 20 |
| α-helix | 830-842 | 13 | |
| α-helix | 848-849 | 2 | |
Chain CH: 6 helices, 2 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 26-33 | 8 | |
| α-helix | 58-63 | 6 | |
| α-helix | 75-88 | 14 | |
| α-helix | 92-99 | 8 | |
| α-helix | 103-110 | 8 | |
| α-helix | 118-128 | 11 | |
| β-strand | 131 | 1 | 63 |
| β-strand | 139 | 1 | 63 |
Chain CI: 2 helices, 0 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 52-62 | 11 | |
| α-helix | 70-72 | 3 | |
Chain La: 7 helices, 7 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 3-5 | 3 | |
| α-helix | 7-9 | 3 | |
| α-helix | 42-48 | 7 | |
| β-strand | 72-74 | 3 | 102 |
| α-helix | 78-81 | 4 | |
| α-helix | 84-92 | 9 | |
| β-strand | 97-98 | 2 | 9 |
| β-strand | 100-102 | 3 | 103 |
| α-helix | 103-106 | 4 | |
| β-strand | 110-112 | 3 | 102 |
| β-strand | 123-126 | 4 | 103 |
| β-strand | 128-129 | 2 | 102 |
| α-helix | 131-139 | 9 | |
| β-strand | 143-146 | 4 | 103 |
Chain LA: 11 helices, 20 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 4-5 | 2 | |
| α-helix | 9-11 | 3 | |
| β-strand | 18 | 1 | 108 |
| α-helix | 29-32 | 4 | |
| α-helix | 34-38 | 5 | |
| β-strand | 40-50 | 11 | 109 |
| α-helix | 51 | 1 | |
| β-strand | 57-64 | 8 | 109 |
| β-strand | 71-77 | 7 | 109 |
| β-strand | 79 | 1 | 110 |
| β-strand | 83 | 1 | 29 |
| β-strand | 87-91 | 5 | 109 |
| β-strand | 101-103 | 3 | 109 |
| α-helix | 104-106 | 3 | |
| β-strand | 112-113 | 2 | 111 |
| β-strand | 116 | 1 | 112 |
| β-strand | 126 | 1 | 112 |
| β-strand | 134-140 | 7 | 111 |
| α-helix | 142-144 | 3 | |
| β-strand | 145-149 | 5 | 111 |
| β-strand | 155-159 | 5 | 111 |
| β-strand | 163-165 | 3 | 109 |
| β-strand | 166 | 1 | 111 |
| β-strand | 169 | 1 | 110 |
| α-helix | 174-176 | 3 | |
| α-helix | 182-190 | 9 | |
| β-strand | 193 | 1 | 108 |
| α-helix | 201-203 | 3 | |
| β-strand | 224-225 | 2 | 113 |
| α-helix | 229-230 | 2 | |
| β-strand | 237-238 | 2 | 113 |
Chain Lb: 6 helices, 0 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 12-18 | 7 | |
| α-helix | 22-24 | 3 | |
| α-helix | 37-48 | 12 | |
| α-helix | 54-73 | 20 | |
| α-helix | 92-100 | 9 | |
| α-helix | 106-117 | 12 | |
Chain LB: 17 helices, 20 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 9-10 | 2 | |
| α-helix | 32-35 | 4 | |
| α-helix | 40-41 | 2 | |
| β-strand | 42-43 | 2 | 122 |
| β-strand | 45-59 | 15 | 123 |
| β-strand | 71-80 | 10 | 123 |
| α-helix | 82-83 | 2 | |
| β-strand | 84-95 | 12 | 124 |
| β-strand | 98-106 | 9 | 124 |
| α-helix | 107 | 1 | |
| α-helix | 112-115 | 4 | |
| α-helix | 116-118 | 3 | |
| α-helix | 140-156 | 17 | |
| β-strand | 159-165 | 7 | 124 |
| α-helix | 168-170 | 3 | |
| β-strand | 180-184 | 5 | 124 |
| β-strand | 185-186 | 2 | 122 |
| α-helix | 190-199 | 10 | |
| β-strand | 204-205 | 2 | 124 |
| α-helix | 207-210 | 4 | |
| β-strand | 216-222 | 7 | 123 |
| α-helix | 223-224 | 2 | |
| β-strand | 225-229 | 5 | 125 |
| α-helix | 231-234 | 4 | |
| α-helix | 237-241 | 5 | |
| β-strand | 270-273 | 4 | 125 |
| β-strand | 276-287 | 12 | 123 |
| β-strand | 292-293 | 2 | 126 |
| β-strand | 297-298 | 2 | 126 |
| α-helix | 300-302 | 3 | |
| β-strand | 320 | 1 | 127 |
| β-strand | 324 | 1 | 127 |
| β-strand | 330-334 | 5 | 123 |
| β-strand | 344-349 | 6 | 123 |
| β-strand | 365-368 | 4 | 123 |
| α-helix | 382-389 | 8 | |
| α-helix | 393-402 | 10 | |
Chain Lc: 4 helices, 4 β-strands
| Element | Residues | Length | Sheet |
|---|
| α-helix | 10-21 | 12 | |
| β-strand | 26-29 | 4 | 105 |
| α-helix | 31-40 | 10 | |
| β-strand | 45-49 | 5 | 105 |
| α-helix | 54-67 | 14 | |
| β-strand | 70-74 | 5 | 105 |
| α-helix | 78-85 | 8 | |
| β-strand | 93-98 | 6 | 105 |
71 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 |
|---|
| A/P site tRNA | AP | RNA | 71 | Homo sapiens | |
| Elongation factor 2 | CB | protein | 856 | Homo sapiens | P13639 (AlphaFold model) |
| P/E site tRNA | PE | RNA | 75 | Homo sapiens | |
| Small ribosomal subunit protein uS3 | SD | protein | 227 | Homo sapiens | P23396 (AlphaFold model) |
| 40S ribosomal protein S5 | SF | protein | 189 | Homo sapiens | P46782 (AlphaFold model) |
| 40S ribosomal protein S10 | SK | protein | 98 | Homo sapiens | P46783 (AlphaFold model) |
| Small ribosomal subunit protein eS12 | SM | protein | 122 | Homo sapiens | P25398 |
| 40S ribosomal protein S15 | SP | protein | 127 | Homo sapiens | P62841 |
| Small ribosomal subunit protein uS9 | SQ | protein | 144 | Homo sapiens | P62249 |
| Small ribosomal subunit protein eS17 | SR | protein | 135 | Homo sapiens | P08708 |
| 40S ribosomal protein S18 | SS | protein | 145 | Homo sapiens | P62269 |
| 40S ribosomal protein S19 | ST | protein | 143 | Homo sapiens | P39019 |
73 more molecules are not listed.
Sequence of entity 1 (AP), FASTA
>9P73_1 A/P site tRNA (chains AP)
GUCUCCGUAGUGUAGGGUAUCACGUUCGCCUAACACGCGAAAGGUCCUCGGUUCGAAACC
GGGCGGAAACA
Sequence of entity 2 (CB), FASTA
>9P73_2 Elongation factor 2 (chains CB)
NFTVDQIRAIMDKKANIRNMSVIAHVDHGKSTLTDSLVCKAGIIASARAGETRFTDTRKD
EQERCITIKSTAISLFYELSENDLNFIKQSKDGAGFLINLIDSPGHVDFSSEVTAALRVT
DGALVVVDCVSGVCVQTETVLRQAIAERIKPVLMMNKMDRALLELQLEPEELYQTFQRIV
ENVNVIISTYGEGESGPMGNIMIDPVLGTVGFGSGLHGWAFTLKQFAEMYVAKFAAKGEG
QLGPAERAKKVEDMMKKLWGDRYFDPANGKFSKSATSPEGKKLPRTFCQLILDPIFKVFD
AIMNFKKEETAKLIEKLDIKLDSEDKDKEGKPLLKAVMRRWLPAGDALLQMITIHLPSPV
TAQKYRCELLYEGPPDDEAAMGIKSCDPKGPLMMYISKMVPTSDKGRFYAFGRVFSGLVS
TGLKVRIMGPNYTPGKKEDLYLKPIQRTILMMGRYVEPIEDVPCGNIVGLVGVDQFLVKT
GTITTFEHAHNMRVMKFSVSPVVRVAVEAKNPADLPKLVEGLKRLAKSDPMVQCIIEESG
EHIIAGAGELHLEICLKDLEEDHACIPIKKSDPVVSYRETVSEESNVLCLSKSPNKHNRL
YMKARPFPDGLAEDIDKGEVSARQELKQRARYLAEKYEWDVAEARKIWCFGPDGTGPNIL
TDITKGVQYLNEIKDSVVAGFQWATKEGALCEENMRGVRFDVHDVTLHADAIHRGGGQII
PTARRCLYASVLTAQPRLMEPIYLVEIQCPEQVVGGIYGVLNRKRGHVFEESQVAGTPMF
VVKAYLPVNESFGFTADLRSNTGGQAFPQCVFDHWQILPGDPFDNSSRPSQVVAETRKRK
GLKEGIPALDNFLDKL
Sequence of entity 3 (PE), FASTA
>9P73_3 P/E site tRNA (chains PE)
GCCCGGAUAGCUCAGCUGUAGAGCAUCAGACUUUUAAUCUGAGGGUCCAGGGUUCAAGUC
CCUGUUCGGGCGCCA
Sequence of entity 4 (SD), FASTA
>9P73_4 Small ribosomal subunit protein uS3 (chains SD)
MAVQISKKRKFVADGIFKAELNEFLTRELAEDGYSGVEVRVTPTRTEIIILATRTQNVLG
EKGRRIRELTAVVQKRFGFPEGSVELYAEKVATRGLCAIAQAESLRYKLLGGLAVRRACY
GVLRFIMESGAKGCEVVVSGKLRGQRAKSMKFVDGLMIHSGDPVNYYVDTAVRHVLLRQG
VLGIKVKIMLPWDPTGKIGPKKPLPDHVSIVEPKDEILPTTPISEQK
Sequence of entity 5 (SF), FASTA
>9P73_5 40S ribosomal protein S5 (chains SF)
DIKLFGKWSTDDVQINDISLQDYIAVKEKYAKYLPHSAGRYAAKRFRKAQCPIVERLTNS
MMMHGRNNGKKLMTVRIVKHAFEIIHLLTGENPLQVLVNAIINSGPREDSTRIGRAGTVR
RQAVDVSPLRRVNQAIWLLCTGAREAAFRNIKTIAECLADELINAAKGSSNSYAIKKKDE
LERVAKSNR
Sequence of entity 6 (SK), FASTA
>9P73_6 40S ribosomal protein S10 (chains SK)
MLMPKKNRIAIYELLFKEGVMVAKKDVHMPKHPELADKNVPNLHVMKAMQSLKSRGYVKE
QFAWRHFYWYLTNEGIQYLRDYLHLPPEIVPATLRRSR
Sequence of entity 7 (SM), FASTA
>9P73_7 Small ribosomal subunit protein eS12 (chains SM)
VMDVNTALQEVLKTALIHDGLARGIREAAKALDKRQAHLCVLASNCDEPMYVKLVEALCA
EHQINLIKVDDNKKLGEWVGLCKIDREGKPRKVVGCSCVVVKDYGKESQAKDVIEEYFKC
KK
Sequence of entity 8 (SP), FASTA
>9P73_8 40S ribosomal protein S15 (chains SP)
RTFRKFTYRGVDLDQLLDMSYEQLMQLYSARQRRRLNRGLRRKQHSLLKRLRKAKKEAPP
MEKPEVVKTHLRDMIILPEMVGSMVGVYNGKTFNQVEIKPEMIGHYLGEFSITYKPVKHG
RPGIGAT
Sequence of entity 9 (SQ), FASTA
>9P73_9 Small ribosomal subunit protein uS9 (chains SQ)
SKGPLQSVQVFGRKKTATAVAHCKRGNGLIKVNGRPLEMIEPRTLQYKLLEPVLLLGKER
FAGVDIRVRVKGGGHVAQIYAIRQSISKALVAYYQKYVDEASKKEIKDILIQYDRTLLVA
DPRRCESKKFGGPGARARYQKSYR
Sequence of entity 10 (SR), FASTA
>9P73_10 Small ribosomal subunit protein eS17 (chains SR)
MGRVRTKTVKKAARVIIEKYYTRLGNDFHTNKRVCEEIAIIPSKKLRNKIAGYVTHLMKR
IQRGPVRGISIKLQEEERERRDNYVPEVSALDQEIIEVDPDTKEMLKLLDFGSLSNLQVT
QPTVGMNFKTPRGPV
Sequence of entity 11 (SS), FASTA
>9P73_11 40S ribosomal protein S18 (chains SS)
MSLVIPEKFQHILRVLNTNIDGRRKIAFAITAIKGVGRRYAHVVLRKADIDLTKRAGELT
EDEVERVITIMQNPRQYKIPDWFLNRQKDVKDGKYSQVLANGLDNKLREDLERLKKIRAH
RGLRHFWGLRVRGQHTKTTGRRGRT
Sequence of entity 12 (ST), FASTA
>9P73_12 40S ribosomal protein S19 (chains ST)
PGVTVKDVNQQEFVRALAAFLKKSGKLKVPEWVDTVKLAKHKELAPYDENWFYTRAASTA
RHLYLRGGAGVGSMTKIYGGRQRNGVMPSHFSRGSKSVARRVLQALEGLKMVEKDQDGGR
KLTPQGQRDLDRIAGQVAAANKK
Ligands and cofactors
| ID | Name | Formula | Copies |
|---|
| MG | Magnesium ion | Mg | 218 |
| ZN | Zinc ion | Zn | 6 |
| SPM | Spermine | C10 H26 N4 | 7 |
| SPD | Spermidine | C7 H19 N3 | 10 |
Primary citation
Visualizing the translation landscape in human cells at high resolution. Zheng, W., Zhang, Y., Wang, J. et al. Nat Commun (2025) 16:10757-10757. DOI 10.1038/s41467-025-65795-9 · PubMed
Other PDB entries of the same protein (UniProt P13639 (AlphaFold model), which also has an AlphaFold model), best resolution first:
- 29IZ 2.08 Å, Dormant 80S:eEF2:SERBP1:tRNA complex from human cells treated with toxin NT-2
- 11IQ 2.4 Å, Rabbit 80S with eEF2,eIF5A and SERBP1
- 8XSX 2.4 Å, Cryo-EM structure of the human 80S ribosome with Tigecycline, E-tRNA, SERBP1 and eEF2
- 9P7D 2.57 Å, In situ human Hibernating class4 80S ribosome
- 9P7E 2.59 Å, In situ human Hibernating class5 80S ribosome
- 9T6D 2.67 Å, NAA40-NAC bound human 80S (combined translation states)
- 9P9I 2.77 Å, In situ human unrotated hibernating without CCDC124 state 80S ribosome
- 9M0P 2.78 Å, Cryo-EM structure of human 80S ribosome in complex with montanine
- 9P7C 2.78 Å, In situ human Hibernating class3 80S ribosome
- 9P7A 2.81 Å, In situ human hibernating class2 80S ribosome
- 9B0P 2.82 Å, In situ human Hibernating class1 80S ribosome
- 9P9H 2.84 Å, In situ human unrotated hibernating with CCDC124 state 80S ribosome
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