9RPV: ZAK-bound human disome

Structure of the ZAK-bound human disome. Determined by electron microscopy at 2.35 Å resolution. Released 10 Dec 2025.

Method
Electron microscopy
Resolution
2.35 Å
Organism
Homo sapiens
Chains
168
Atoms
428,966
Mol. weight
7974.13 kDa
Ligands
MG, ZN, ANM
Released
10 Dec 2025

Explore 9RPV in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9RPV contains 1,138 α-helices and 1,091 β-strands across 154 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 E4: 9 helices, 1 β-strand

ElementResiduesLengthSheet
α-helix101
β-strand11172
α-helix121
α-helix25-317
α-helix58-636
α-helix72-743
α-helix75-8612
α-helix92-987
α-helix103-1119
α-helix118-12811
Chain La: 9 helices, 7 β-strands
ElementResiduesLengthSheet
α-helix3-53
α-helix7-104
α-helix42-487
β-strand72-743124
α-helix75-773
α-helix79-813
α-helix84-929
β-strand981285
β-strand100-1023125
α-helix103-1064
β-strand110-1123124
α-helix116-1183
β-strand123-1264125
β-strand128-1292124
α-helix131-14010
β-strand143-1464125
Chain LA: 13 helices, 18 β-strands
ElementResiduesLengthSheet
α-helix3-53
α-helix10-123
α-helix14-163
β-strand181137
α-helix30-323
α-helix34-385
β-strand41-499138
β-strand58-647138
β-strand71-777138
α-helix781
β-strand791139
β-strand831140
β-strand87-904138
β-strand101-1033138
α-helix104-1063
α-helix1081
β-strand112-1132141
β-strand134-1407141
α-helix142-1443
β-strand145-1495141
β-strand155-1595141
β-strand163-1653138
β-strand1661141
β-strand1691139
α-helix174-1763
α-helix182-1909
β-strand1931137
α-helix201-2033
β-strand224-2252142
α-helix229-2302
β-strand237-2382142
Chain Lb: 4 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix12-154
α-helix22-243
α-helix37-4812
α-helix54-629
Chain LB: 19 helices, 18 β-strands
ElementResiduesLengthSheet
α-helix9-102
α-helix14-163
α-helix20-223
α-helix32-354
α-helix40-412
β-strand421156
β-strand45-6016157
β-strand70-8011157
β-strand84-9512158
β-strand98-1069158
α-helix112-1154
α-helix116-1183
α-helix131-1344
α-helix135-1373
α-helix140-15617
β-strand159-1657158
α-helix168-1703
β-strand180-1845158
β-strand1861156
α-helix190-19910
β-strand204-2052158
α-helix207-2093
β-strand216-2227157
β-strand227-2293159
α-helix231-2344
α-helix237-2415
β-strand270-2723159
β-strand276-28712157
α-helix300-3023
β-strand3201160
β-strand3241160
β-strand330-3334157
β-strand344-3496157
α-helix361-3633
β-strand365-3684157
α-helix382-3898
α-helix393-3986
Chain Lc: 4 helices, 4 β-strands
ElementResiduesLengthSheet
α-helix15-217
β-strand27-293161
α-helix32-387
β-strand45-484161
α-helix54-6613
β-strand70-734161
α-helix78-847
β-strand93-964161
Chain LC: 25 helices, 12 β-strands
ElementResiduesLengthSheet
β-strand7-115178
β-strand17-237178
α-helix24-252
α-helix26-294
α-helix34-4512
α-helix46-483
α-helix49-513
β-strand64-663179
β-strand77-793179
β-strand901179
α-helix100-1012
α-helix117-13115
α-helix134-1407
β-strand152-1532178
α-helix156-1605
α-helix164-17310
α-helix178-1858
β-strand188-1892180
α-helix1901
α-helix193-1964
β-strand201-2022180
β-strand208-2114178
α-helix217-2215
β-strand228-2314178
β-strand2341178
α-helix237-2404
β-strand250-2523178
α-helix254-2585
α-helix260-2634
α-helix279-2824
α-helix290-2934
α-helix296-3005
α-helix302-3054
α-helix311-3133
α-helix316-3172
α-helix322-3287
α-helix332-36231
Chain Ld: 4 helices, 7 β-strands
ElementResiduesLengthSheet
β-strand231181
β-strand26-272182
α-helix38-403
α-helix41-5717
β-strand62-643183
α-helix66-727
β-strand84-852182
β-strand86-927183
β-strand102-1087183
β-strand1211181
α-helix122-1232

145 more chain groups are not listed. Open the entry in the viewer and use the sequence panel to see them.

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
mRNA of the collided 80SA4RNA14Homo sapiens
mRNA of the stalled 80SA5RNA8Homo sapiens
P/E tRNA on the collided 80SB4RNA150Homo sapiens
P-site tRNA of the stalled 80SB5RNA77Homo sapiens
A/P tRNA on the collided 80SD4RNA152Homo sapiens
Endothelial differentiation-related factor 1E4protein148Homo sapiensO60869 (AlphaFold model)
E-site tRNA of the stalled 80SE5RNA75Homo sapiens
28S rRNA of the collided 80SL5RNA5070Homo sapiens
5S rRNAL7, M7RNA120Homo sapiens
5.8S rRNA of the collided 80SL8RNA156Homo sapiens
60S ribosomal protein L8LAprotein257Homo sapiensP62917 (AlphaFold model)
60S ribosomal protein L3LBprotein403Homo sapiensP39023 (AlphaFold model)

89 more molecules are not listed.

Sequence of entity 1 (A4), FASTA
>9RPV_1 mRNA of the collided 80S (chains A4)
AUUUUAUGUUCUUU
Sequence of entity 2 (A5), FASTA
>9RPV_2 mRNA of the stalled 80S (chains A5)
AACGUAUG
Sequence of entity 3 (B4), FASTA
>9RPV_3 P/E tRNA on the collided 80S (chains B4)
AGCAGAGUGGCGCAGCGGAAGCGUGCUGGGCCCAUAACCCAGAGGUCGAUGGAUCGAAAC
CAUCCUCUGCUACCAAGCAGAGUGGCGCAGCGGAAGCGUGCUGGGCCCAUAACCCAGAGG
UCGAUGGAUCGAAACCAUCCUCUGCUACCA
Sequence of entity 4 (B5), FASTA
>9RPV_4 P-site tRNA of the stalled 80S (chains B5)
CGCGGGGUGGAGCAGCCCGGUAGCUCGUCGGGCUCAUAACCCGAAGGUCGUCGGUUCAAA
UCCGGCCCCCGCAACCA
Sequence of entity 5 (D4), FASTA
>9RPV_5 A/P tRNA on the collided 80S (chains D4)
GCCCGGAUAGCUCAGUCGGUAGAGCAGGGGAUUGAAAAUCCCCGUGUCCUUGGUUCGAUU
CCGAGUCCGGGCACCAGCCCGGAUAGCUCAGUCGGUAGAGCAGGGGAUUGAAAAUCCCCG
UGUCCUUGGUUCGAUUCCGAGUCCGGGCACCA
Sequence of entity 6 (E4), FASTA
>9RPV_6 Endothelial differentiation-related factor 1 (chains E4)
MAESDWDTVTVLRKKGPTAAQAKSKQAILAAQRRGEDVETSKKWAAGQNKQHSITKNTAK
LDRETEELHHDRVTLEVGKVIQQGRQSKGLTQKDLATKINEKPQVIADYESGRAIPNNQV
LGKIERAIGLKLRGKDIGKPIEKGPRAK
Sequence of entity 7 (E5), FASTA
>9RPV_7 E-site tRNA of the stalled 80S (chains E5)
UCCGAUAUAGCGUAACGGCUAUCACAUCACGCUUUCACCGUGGAGACCGGGGUUCGACUC
CCCGUAUCGGAGCCA
Sequence of entity 8 (L5), FASTA
>9RPV_8 28S rRNA of the collided 80S (chains L5)
CGCGACCUCAGAUCAGACGUGGCGACCCGCUGAAUUUAAGCAUAUUAGUCAGCGGAGGAG
AAGAAACUAACCAGGAUUCCCUCAGUAACGGCGAGUGAACAGGGAAGAGCCCAGCGCCGA
AUCCCCGCCCCGCGGCGGGGCGCGGGACAUGUGGCGUACGGAAGACCCGCUCCCCGGCGC
CGCUCGUGGGGGGCCCAAGUCCUUCUGAUCGAGGCCCAGCCCGUGGACGGUGUGAGGCCG
GUAGCGGCCCCCGGCGCGCCGGGCCCGGGUCUUCCCGGAGUCGGGUUGCUUGGGAAUGCA
GCCCAAAGCGGGUGGUAAACUCCAUCUAAGGCUAAAUACCGGCACGAGACCGAUAGUCAA
CAAGUACCGUAAGGGAAAGUUGAAAAGAACUUUGAAGAGAGAGUUCAAGAGGGCGUGAAA
CCGUUAAGAGGUAAACGGGUGGGGUCCGCGCAGUCCGCCCGGAGGAUUCAACCCGGCGGC
GGGUCCGGCCGUGUCGGCGGCCCGGCGGAUCUUUCCCGCCCCCCGUUCCUCCCGACCCCU
CCACCCGCCCUCCCUUCCCCCGCCGCCCCUCCUCCUCCUCCCCGGAGGGGGCGGGCUCCG
GCGGGUGCGGGGGUGGGCGGGCGGGGCCGGGGGUGGGGUCGGCGGGGGACCGUCCCCCGA
CCGGCGACCGGCCGCCGCCGGGCGCAUUUCCACCGCGGCGGUGCGCCGCGACCGGCUCCG
GGACGGCUGGGAAGGCCCGGCGGGGAAGGUGGCUCGGGGGGCCCCGUCCGUCCGUCCGUC
CGUCCUCCUCCUCCCCCGUCUCCGCCCCCCGGCCCCGCGUCCUCCCUCGGGAGGGCGCGC
GGGUCGGGGCGGCGGCGGCGGCGGCGGUGGCGGCGGCGGCGGCGGCGGCGGGACCGAAAC
CCCCCCCGAGUGUUACAGCCCCCCCGGCAGCAGCACUCGCCGAAUCCCGGGGCCGAGGGA
GCGAGACCCGUCGCCGCGCUCUCCCCCCUCCCGGCGCCCACCCCCGCGGGGAAUCCCCCG
CGAGGGGGGUCUCCCCCGCGGGGGCGCGCCGGCGUCUCCUCGUGGGGGGGCCGGGCCACC
CCUCCCACGGCGCGACCGCUCUCCCACCCCUCCUCCCCGCGCCCCCGCCCCGGCGACGGG
GGGGGUGCCGCGCGCGGGUCGGGGGGCGGGGCGGACUGUCCCCAGUGCGCCCCGGGCGGG
UCGCGCCGUCGGGCCCGGGGGAGGUUCUCUCGGGGCCACGCGCGCGUCCCCCGAAGAGGG
GGACGGCGGAGCGAGCGCACGGGGUCGGCGGCGACGUCGGCUACCCACCCGACCCGUCUU
GAAACACGGACCAAGGAGUCUAACACGUGCGCGAGUCGGGGGCUCGCACGAAAGCCGCCG
UGGCGCAAUGAAGGUGAAGGCCGGCGCGCUCGCCGGCCGAGGUGGGAUCCCGAGGCCUCU
CCAGUCCGCCGAGGGCGCACCACCGGCCCGUCUCGCCCGCCGCGCCGGGGAGGUGGAGCA
CGAGCGCACGUGUUAGGACCCGAAAGAUGGUGAACUAUGCCUGGGCAGGGCGAAGCCAGA
GGAAACUCUGGUGGAGGUCCGUAGCGGUCCUGACGUGCAAAUCGGUCGUCCGACCUGGGU
AUAGGGGCGAAAGACUAAUCGAACCAUCUAGUAGCUGGUUCCCUCCGAAGUUUCCCUCAG
GAUAGCUGGCGCUCUCGCAGACCCGACGCACCCCCGCCACGCAGUUUUAUCCGGUAAAGC
GAAUGAUUAGAGGUCUUGGGGCCGAAACGAUCUCAACCUAUUCUCAAACUUUAAAUGGGU
AAGAAGCCCGGCUCGCUGGCGUGGAGCCGGGCGUGGAAUGCGAGUGCCUAGUGGGCCACU
UUUGGUAAGCAGAACUGGCGCUGCGGGAUGAACCGAACGCCGGGUUAAGGCGCCCGAUGC
CGACGCUCAUCAGACCCCAGAAAAGGUGUUGGUUGAUAUAGACAGCAGGACGGUGGCCAU
GGAAGUCGGAAUCCGCUAAGGAGUGUGUAACAACUCACCUGCCGAAUCAACUAGCCCUGA
AAAUGGAUGGCGCUGGAGCGUCGGGCCCAUACCCGGCCGUCGCCGGCAGUCGAGAGUGGA
CGGGAGCGGCGGCGGCGGCGCGCGCGCGCGCGCGUGUGGUGUGCGUCGGAGGGCGGCGGC
GGCGGCGGCGGCGGGGGUGUGGGGUCCUUCCCCCGCCCCCCCCCCCACGCCUCCUCCCCU
CCUCCCGCCCACGCCCCGCUCCCCGCCCCCGGAGCCCCGCGGACGCUACGCCGCGACGAG
UAGGAGGGCCGCUGCGGUGAGCCUUGAAGCCUAGGGCGCGGGCCCGGGUGGAGCCGCCGC
AGGUGCAGAUCUUGGUGGUAGUAGCAAAUAUUCAAACGAGAACUUUGAAGGCCGAAGUGG
AGAAGGGUUCCAUGUGAACAGCAGUUGAACAUGGGUCAGUCGGUCCUGAGAGAUGGGCGA
GCGCCGUUCCGAAGGGACGGGCGAUGGCCUCCGUUGCCCUCGGCCGAUCGAAAGGGAGUC
GGGUUCAGAUCCCCGAAUCCGGAGUGGCGGAGAUGGGCGCCGCGAGGCGUCCAGUGCGGU
AACGCGACCGAUCCCGGAGAAGCCGGCGGGAGCCCCGGGGAGAGUUCUCUUUUCUUUGUG
AAGGGCAGGGCGCCCUGGAAUGGGUUCGCCCCGAGAGAGGGGCCCGUGCCUUGGAAAGCG
UCGCGGUUCCGGCGGCGUCCGGUGAGCUCUCGCUGGCCCUUGAAAAUCCGGGGGAGAGGG
UGUAAAUCUCGCGCCGGGCCGUACCCAUAUCCGCAGCAGGUCUCCAAGGUGAACAGCCUC
UGGCAUGUUGGAACAAUGUAGGUAAGGGAAGUCGGCAAGCCGGAUCCGUAACUUCGGGAU
AAGGAUUGGCUCUAAGGGCUGGGUCGGUCGGGCUGGGGCGCGAAGCGGGGCUGGGCGCGC
GCCGCGGCUGGACGAGGCGCCGCCGCCCCCCCCACGCCCGGGGCACCCCCCUCGCGGCCC
UCCCCCGCCCCACCCCGCGCGCGCCGCUCGCUCCCUCCCCGCCCCGCGCCCUCUCUCUCU
CUCUCUCCCCCGCUCCCCGUCCUCCCCCCUCCCCGGGGGAGCGCCGCGUGGGGGCGGCGG
CGGGGGGAGAAGGGUCGGGGCGGCAGGGGCCGGCGGCGGCCCGCCGCGGGGCCCCGGCGG
CGGGGGCACGGUCCCCCGCGAGGGGGGCCCGGGCACCCGGGGGGCCGGCGGCGGCGGCGA
CUCUGGACGCGAGCCGGGCCCUUCCCGUGGAUCGCCCCAGCUGCGGCGGGCGUCGCGGCC
GCCCCCGGGGAGCCCGGCGGGCGCCGGCGCGCCCCCCCCCCCACCCCACGUCUCGUCGCG
CGCGCGUCCGCUGGGGGCGGGGAGCGGUCGGGCGGCGGCGGUCGGCGGGCGGCGGGGCGG
GGCGGUUCGUCCCCCCGCCCUACCCCCCCGGCCCCGUCCGCCCCCCGUUCCCCCCUCCUC
CUCGGCGCGCGGCGGCGGCGGCGGCAGGCGGCGGAGGGGCCGCGGGCCGGUCCCCCCCGC
CGGGUCCGCCCCCGGGGCCGCGGUUCCGCGCGGCGCCUCGCCUCGGCCGGCGCCUAGCAG
CCGACUUAGAACUGGUGCGGACCAGGGGAAUCCGACUGUUUAAUUAAAACAAAGCAUCGC
GAAGGCCCGCGGCGGGUGUUGACGCGAUGUGAUUUCUGCCCAGUGCUCUGAAUGUCAAAG
UGAAGAAAUUCAAUGAAGCGCGGGUAAACGGCGGGAGUAACUAUGACUCUCUUAAGGUAG
CCAAAUGCCUCGUCAUCUAAUUAGUGACGCGCAUGAAUGGAUGAACGAGAUUCCCACUGU
CCCUACCUACUAUCCAGCGAAACCACAGCCAAGGGAACGGGCUUGGCGGAAUCAGCGGGG
AAAGAAGACCCUGUUGAGCUUGACUCUAGUCUGGCACGGUGAAGAGACAUGAGAGGUGUA
GAAUAAGUGGGAGGCCCCCGGCGCCCCCCCGGUGUCCCCGCGAGGGGCCCGGGGCGGGGU
CCGCCGGCCCUGCGGGCCGCCGGUGAAAUACCACUACUCUGAUCGUUUUUUCACUGACCC
GGUGAGGCGGGGGGGCGAGCCCCGAGGGGCUCUCGCUUCUGGCGCCAAGCGCCCGGCCGC
GCGCCGGCCGGGCGCGACCCGCUCCGGGGACAGUGCCAGGUGGGGAGUUUGACUGGGGCG
GUACACCUGUCAAACGGUAACGCAGGUGUCCUAAGGCGAGCUCAGGGAGGACAGAAACCU
CCCGUGGAGCAGAAGGGCAAAAGCUCGCUUGAUCUUGAUUUUCAGUACGAAUACAGACCG
UGAAAGCGGGGCCUCACGAUCCUUCUGACCUUUUGGGUUUUAAGCAGGAGGUGUCAGAAA
AGUUACCACAGGGAUAACUGGCUUGUGGCGGCCAAGCGUUCAUAGCGACGUCGCUUUUUG
AUCCUUCGAUGUCGGCUCUUCCUAUCAUUGUGAAGCAGAAUUCACCAAGCGUUGGAUUGU
UCACCCACUAAUAGGGAACGUGAGCUGGGUUUAGACCGUCGUGAGACAGGUUAGUUUUAC
CCUACUGAUGAUGUGUUGUUGCCAUGGUAAUCCUGCUCAGUACGAGAGGAACCGCAGGUU
CAGACAUUUGGUGUAUGUGCUUGGCUGAGGAGCCAAUGGGGCGAAGCUACCAUCUGUGGG
AUUAUGACUGAACGCCUCUAAGUCAGAAUCCCGCCCAGGCGGAACGAUACGGCAGCGCCG
CGGAGCCUCGGUUGGCCUCGGAUAGCCGGUCCCCCGCCUGUCCCCGCCGGCGGGCCGCCC
CCCCCCUCCACGCGCCCCGCGCGCGCGGGAGGGCGCGUGCCCCGCCGCGCGCCGGGACCG
GGGUCCGGUGCGGAGUGCCCUUCGUCCUGGGAAACGGGGCGCGGCCGGAGAGGCGGCCGC
CCCCUCGCCCGUCACGCACCGCACGUUCGUGGGGAACCUGGCGCUAAACCAUUCGUAGAC
GACCUGCUUCUGGGUCGGGG
Sequence of entity 9 (L7, M7), FASTA
>9RPV_9 5S rRNA (chains L7, M7)
GUCUACGGCCAUACCACCCUGAACGCGCCCGAUCUCGUCUGAUCUCGGAAGCUAAGCAGG
GUCGGGCCUGGUUAGUACUUGGAUGGGAGACCGCCUGGGAAUACCGGGUGCUGUAGGCUU
Sequence of entity 10 (L8), FASTA
>9RPV_10 5.8S rRNA of the collided 80S (chains L8)
CGACUCUUAGCGGUGGAUCACUCGGCUCGUGCGUCGAUGAAGAACGCAGCUAGCUGCGAG
AAUUAAUGUGAAUUGCAGGACACAUUGAUCAUCGACACUUCGAACGCACUUGCGGCCCCG
GGUUCCUCCCGGGGCUACGCCUGUCUGAGCGUCGCU
Sequence of entity 11 (LA), FASTA
>9RPV_11 60S ribosomal protein L8 (chains LA)
MGRVIRGQRKGAGSVFRAHVKHRKGAARLRAVDFAERHGYIKGIVKDIIHDPGRGAPLAK
VVFRDPYRFKKRTELFIAAEGIHTGQFVYCGKKAQLNIGNVLPVGTMPEGTIVCCLEEKP
GDRGKLARASGNYATVISHNPETKKTRVKLPSGSKKVISSANRAVVGVVAGGGRIDKPIL
KAGRAYHKYKAKRNCWPRVRGVAMNPVEHPFGGGNHQHIGKPSTIRRDAPAGRKVGLIAA
RRTGRLRGTKTVQEKEN
Sequence of entity 12 (LB), FASTA
>9RPV_12 60S ribosomal protein L3 (chains LB)
MSHRKFSAPRHGSLGFLPRKRSSRHRGKVKSFPKDDPSKPVHLTAFLGYKAGMTHIVREV
DRPGSKVNKKEVVEAVTIVETPPMVVVGIVGYVETPRGLRTFKTVFAEHISDECKRRFYK
NWHKSKKKAFTKYCKKWQDEDGKKQLEKDFSSMKKYCQVIRVIAHTQMRLLPLRQKKAHL
MEIQVNGGTVAEKLDWARERLEQQVPVNQVFGQDEMIDVIGVTKGKGYKGVTSRWHTKKL
PRKTHRGLRKVACIGAWHPARVAFSVARAGQKGYHHRTEINKKIYKIGQGYLIKDGKLIK
NNASTDYDLSDKSINPLGGFVHYGEVTNDFVMLKGCVVGTKKRVLTLRKSLLVQTKRRAL
EKIDLKFIDTTSKFGHGRFQTMEEKKAFMGPLKKDRIAKEEGA

Ligands and cofactors

IDNameFormulaCopies
MGMagnesium ionMg830
ZNZinc ionZn7
ANMAnisomycinC14 H19 N O41

Primary citation

ZAK activation at the collided ribosome. Huso, V.L., Niu, S., Catipovic, M.A. et al. Nature (2026) 649:1051-1060. DOI 10.1038/s41586-025-09772-8 · PubMed

Other PDB entries of the same protein (UniProt O60869 (AlphaFold model), which also has an AlphaFold model), best resolution first:

Browse structure collections

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