28LN: XBP1u-stalled RPL4 RNC

XBP1u-stalled RPL4 RNC in complex with NAC. Determined by electron microscopy at 2.7 Å resolution. Released 10 Jun 2026.

Method
Electron microscopy
Resolution
2.7 Å
Organism
Homo sapiens
Chains
86
Atoms
220,645
Mol. weight
3945.08 kDa
Ligands
MG, ZN
Released
10 Jun 2026

Explore 28LN in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

28LN contains 593 α-helices and 574 β-strands across 80 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 Aa: 5 helices, 5 β-strands

ElementResiduesLengthSheet
β-strand20-2238
β-strand29-3138
β-strand36-4389
α-helix47-5610
β-strand571111
α-helix62-654
β-strand66-7389
α-helix75-806
α-helix89-924
α-helix96-1005
Chain AA: 13 helices, 8 β-strands
ElementResiduesLengthSheet
α-helix10-112
α-helix12-2110
β-strand2516
β-strand38-4146
α-helix461
β-strand47-4936
α-helix501
α-helix51-6717
α-helix71-733
β-strand74-7857
α-helix81-833
α-helix84-9411
β-strand97-10047
α-helix102-1043
β-strand121-12447
α-helix131-1377
β-strand144-14857
β-strand159-16247
α-helix168-18619
α-helix199-2024
α-helix207-21610
Chain Ab: 3 helices, 7 β-strands
ElementResiduesLengthSheet
α-helix12-176
β-strand32-36514
β-strand43-47514
β-strand55115
β-strand62115
β-strand64-65216
α-helix711
β-strand72-73216
α-helix741
β-strand78-81414
Chain AB: 10 helices, 12 β-strands
ElementResiduesLengthSheet
α-helix24-274
β-strand28-34711
β-strand43-50811
α-helix51-522
β-strand53112
β-strand56112
α-helix58-625
β-strand66-70511
α-helix71-744
β-strand82-921111
β-strand95-1051111
α-helix107-1137
β-strand118113
β-strand120-128911
β-strand134-1431011
α-helix144-1463
β-strand154113
α-helix158-17720
α-helix181-1888
α-helix192-2009
β-strand207-2171111
α-helix225-2328
Chain Ac: 1 helix, 4 β-strands
ElementResiduesLengthSheet
β-strand10-211221
β-strand27-34821
β-strand41-47721
β-strand55-58421
α-helix64-674
Chain AC: 11 helices, 10 β-strands
ElementResiduesLengthSheet
α-helix65-717
α-helix79-846
α-helix91-988
α-helix99-1013
β-strand103-1161418
β-strand119-1311318
β-strand136-144918
α-helix147-16014
β-strand163-164218
β-strand168119
β-strand179119
β-strand184-188520
β-strand191-196620
β-strand204-205220
α-helix208-21710
β-strand222-227620
α-helix233-24917
α-helix253-2553
α-helix258-2592
α-helix265-2684
α-helix270-2756
Chain Ad: 4 helices, 3 β-strands
ElementResiduesLengthSheet
α-helix3-64
β-strand30-31230
α-helix33-353
β-strand38-39230
α-helix40-456
α-helix47-504
β-strand531154
Chain AD: 11 helices, 9 β-strands
ElementResiduesLengthSheet
α-helix7-2822
β-strand34-40726
β-strand46-52726
α-helix55-595
α-helix61-633
α-helix64-7714
α-helix79-802
β-strand84-90726
α-helix98-11114
α-helix115-12915
β-strand133-140827
β-strand148-155827
α-helix164-1674
β-strand168-176927
β-strand181-189927
α-helix190-1923
β-strand209-210228
α-helix211-2144
α-helix216-2194
β-strand225129

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

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
Large ribosomal subunit protein uL4,X-box-binding protein 1, luminal form,X-box-binding protein 1,…B1protein184Homo sapiensP17861 (AlphaFold model)
tRNA-Met-CATB2RNA76Homo sapiens
mRNA fragment (6-MER)B4RNA6Homo sapiens
28S rRNAB5RNA5069Homo sapiens
5S rRNAB7RNA120Homo sapiens
5.8S rRNAB8RNA157Homo sapiens
Large ribosomal subunit protein uL2BAprotein257Homo sapiensP62917 (AlphaFold model)
Large ribosomal subunit protein uL3BBprotein403Homo sapiensP39023 (AlphaFold model)
60S ribosomal protein L4BCprotein427Homo sapiensP36578 (AlphaFold model)
60S ribosomal protein L5BDprotein297Homo sapiensP46777
Large ribosomal subunit protein eL6BEprotein288Homo sapiensQ02878
Large ribosomal subunit protein uL30BFprotein248Homo sapiensP18124

74 more molecules are not listed.

Sequence of entity 1 (B1), FASTA
>28LN_1 Large ribosomal subunit protein uL4,X-box-binding protein 1, luminal form,X-box-binding protein 1, luminal form,X-box-binding protein 1, luminal form (chains B1)
MACARPLISVYSEKGESSGKNVTLPAVFKAPIRPDIVNFVHTNLRKNNRQPYAVSELAGH
QTSAESWGTGRAVARIPRVRGGGTHRSGQGAFGNMCRGGRMFAPTKTWRRWHRRVNTTQK
RYAICSALAASALPALVMSKGHRIEEVPELPLVVEDKVQKDPVPYQPPFLCQWGRHQCAW
KPLM
Sequence of entity 2 (B2), FASTA
>28LN_2 tRNA-Met-CAT (chains B2)
GCCUCGUUAGCGCAGUAGGUAGCGCGUCAGUCUCAUAAUCUGAAGGUCGUGAGUUCGAUC
CUCACACGGGGCACCA
Sequence of entity 3 (B4), FASTA
>28LN_3 mRNA fragment (6-MER) (chains B4)
UUAAUG
Sequence of entity 4 (B5), FASTA
>28LN_4 28S rRNA (chains B5)
CGCGACCUCAGAUCAGACGUGGCGACCCGCUGAAUUUAAGCAUAUUAGUCAGCGGAGGAG
AAGAAACUAACCAGGAUUCCCUCAGUAACGGCGAGUGAACAGGGAAGAGCCCAGCGCCGA
AUCCCCGCCCCGCGGCGGGGCGCGGGACAUGUGGCGUACGGAAGACCCGCUCCCCGGCGC
CGCUCGUGGGGGGCCCAAGUCCUUCUGAUCGAGGCCCAGCCCGUGGACGGUGUGAGGCCG
GUAGCGGCCCCCGGCGCGCCGGGCCCGGGUCUUCCCGGAGUCGGGUUGCUUGGGAAUGCA
GCCCAAAGCGGGUGGUAAACUCCAUCUAAGGCUAAAUACCGGCACGAGACCGAUAGUCAA
CAAGUACCGUAAGGGAAAGUUGAAAAGAACUUUGAAGAGAGAGUUCAAGAGGGCGUGAAA
CCGUUAAGAGGUAAACGGGUGGGGUCCGCGCAGUCCGCCCGGAGGAUUCAACCCGGCGGC
GGGUCCGGCCGUGUCGGCGGCCCGGCGGAUCUUUCCCGCCCCCCGUUCCUCCCGACCCCU
CCACCCGCCCUCCCUUCCCCCGCCGCCCCUCCUCCUCCUCCCCGGAGGGGGCGGGCUCCG
GCGGGUGCGGGGGUGGGCGGGCGGGGCCGGGGGUGGGGUCGGCGGGGGACCGUCCCCCGA
CCGGCGACCGGCCGCCGCCGGGCGCAUUUCCACCGCGGCGGUGCGCCGCGACCGGCUCCG
GGACGGCUGGGAAGGCCCGGCGGGGAAGGUGGCUCGGGGGGCCCCGUCCGUCCGUCCGUC
CGUCCUCCUCCUCCCCCGUCUCCGCCCCCCGGCCCCGCGUCCUCCCUCGGGAGGGCGCGC
GGGUCGGGGCGGCGGCGGCGGCGGCGGUGGCGGCGGCGGCGGCGGCGGCGGGACCGAAAC
CCCCCCGAGUGUUACAGCCCCCCCGGCAGCAGCACUCGCCGAAUCCCGGGGCCGAGGGAG
CGAGACCCGUCGCCGCGCUCUCCCCCCUCCCGGCGCCCACCCCCGCGGGGAAUCCCCCGC
GAGGGGGGUCUCCCCCGCGGGGGCGCGCCGGCGUCUCCUCGUGGGGGGGCCGGGCCACCC
CUCCCACGGCGCGACCGCUCUCCCACCCCUCCUCCCCGCGCCCCCGCCCCGGCGACGGGG
GGGGUGCCGCGCGCGGGUCGGGGGGCGGGGCGGACUGUCCCCAGUGCGCCCCGGGCGGGU
CGCGCCGUCGGGCCCGGGGGAGGUUCUCUCGGGGCCACGCGCGCGUCCCCCGAAGAGGGG
GACGGCGGAGCGAGCGCACGGGGUCGGCGGCGACGUCGGCUACCCACCCGACCCGUCUUG
AAACACGGACCAAGGAGUCUAACACGUGCGCGAGUCGGGGGCUCGCACGAAAGCCGCCGU
GGCGCAAUGAAGGUGAAGGCCGGCGCGCUCGCCGGCCGAGGUGGGAUCCCGAGGCCUCUC
CAGUCCGCCGAGGGUGCACCACCGGCCCGUCUCGCCCGCCGCGCCGGGGAGGUGGAGCAC
GAGCGCACGUGUUAGGACCCGAAAGAUGGUGAACUAUGCCUGGGCAGGGCGAAGCCAGAG
GAAACUCUGGUGGAGGUCCGUAGCGGUCCUGACGUGCAAAUCGGUCGUCCGACCUGGGUA
UAGGGGCGAAAGACUAAUCGAACCAUCUAGUAGCUGGUUCCCUCCGAAGUUUCCCUCAGG
AUAGCUGGCGCUCUCGCAGACCCGACGCACCCCCGCCACGCAGUUUUAUCCGGUAAAGCG
AAUGAUUAGAGGUCUUGGGGCCGAAACGAUCUCAACCUAUUCUCAAACUUUAAAUGGGUA
AGAAGCCCGGCUCGCUGGCGUGGAGCCGGGCGUGGAAUGCGAGUGCCUAGUGGGCCACUU
UUGGUAAGCAGAACUGGCGCUGCGGGAUGAACCGAACGCCGGGUUAAGGCGCCCGAUGCC
GACGCUCAUCAGACCCCAGAAAAGGUGUUGGUUGAUAUAGACAGCAGGACGGUGGCCAUG
GAAGUCGGAAUCCGCUAAGGAGUGUGUAACAACUCACCUGCCGAAUCAACUAGCCCUGAA
AAUGGAUGGCGCUGGAGCGUCGGGCCCAUACCCGGCCGUCGCCGGCAGUCGAGAGUGGAC
GGGAGCGGCGGGGGCGGCGCGCGCGCGCGCGCGUGUGGUGUGCGUCGGAGGGCGGCGGCG
GCGGCGGCGGCGGGGGUGUGGGGUCCUUCCCCCGCCCCCCCCCCCACGCCUCCUCCCCUC
CUCCCGCCCACGCCCCGCUCCCCGCCCCCGGAGCCCCGCGGACGCUACGCCGCGACGAGU
AGGAGGGCCGCUGCGGUGAGCCUUGAAGCCUAGGGCGCGGGCCCGGGUGGAGCCGCCGCA
GGUGCAGAUCUUGGUGGUAGUAGCAAAUAUUCAAACGAGAACUUUGAAGGCCGAAGUGGA
GAAGGGUUCCAUGUGAACAGCAGUUGAACAUGGGUCAGUCGGUCCUGAGAGAUGGGCGAG
CGCCGUUCCGAAGGGACGGGCGAUGGCCUCCGUUGCCCUCGGCCGAUCGAAAGGGAGUCG
GGUUCAGAUCCCCGAAUCCGGAGUGGCGGAGAUGGGCGCCGCGAGGCGUCCAGUGCGGUA
ACGCGACCGAUCCCGGAGAAGCCGGCGGGAGCCCCGGGGAGAGUUCUCUUUUCUUUGUGA
AGGGCAGGGCGCCCUGGAAUGGGUUCGCCCCGAGAGAGGGGCCCGUGCCUUGGAAAGCGU
CGCGGUUCCGGCGGCGUCCGGUGAGCUCUCGCUGGCCCUUGAAAAUCCGGGGGAGAGGGU
GUAAAUCUCGCGCCGGGCCGUACCCAUAUCCGCAGCAGGUCUCCAAGGUGAACAGCCUCU
GGCAUGUUGGAACAAUGUAGGUAAGGGAAGUCGGCAAGCCGGAUCCGUAACUUCGGGAUA
AGGAUUGGCUCUAAGGGCUGGGUCGGUCGGGCUGGGGCGCGAAGCGGGGCUGGGCGCGCG
CCGCGGCUGGACGAGGCGCCGCCGCCCCCCCCACGCCCGGGGCACCCCCCUCGCGGCCCU
CCCCCGCCCCACCCCGCGCGCGCCGCUCGCUCCCUCCCCGCCCCGCGCCCUCUCUCUCUC
UCUCUCCCCCGCUCCCCGUCCUCCCCCCUCCCCGGGGGAGCGCCGCGUGGGGGCGGCGGC
GGGGGGAGAAGGGUCGGGGCGGCAGGGGCCGGCGGCGGCCCGCCGCGGGGCCCCGGCGGC
GGGGGCACGGUCCCCCGCGAGGGGGGCCCGGGCACCCGGGGGGCCGGCGGCGGCGGCGAC
UCUGGACGCGAGCCGGGCCCUUCCCGUGGAUCGCCCCAGCUGCGGCGGGCGUCGCGGCCG
CCCCCGGGGAGCCCGGCGGGCGCCGGCGCGCCCCCCCCCCCACCCCACGUCUCGUCGCGC
GCGCGUCCGCUGGGGGCGGGGAGCGGUCGGGCGGCGGCGGUCGGCGGGCGGCGGGGCGGG
GCGGUUCGUCCCCCCGCCCUACCCCCCCGGCCCCGUCCGCCCCCCGUUCCCCCCUCCUCC
UCGGCGCGCGGCGGCGGCGGCGGCAGGCGGCGGAGGGGCCGCGGGCCGGUCCCCCCCGCC
GGGUCCGCCCCCGGGGCCGCGGUUCCGCGCGGCGCCUCGCCUCGGCCGGCGCCUAGCAGC
CGACUUAGAACUGGUGCGGACCAGGGGAAUCCGACUGUUUAAUUAAAACAAAGCAUCGCG
AAGGCCCGCGGCGGGUGUUGACGCGAUGUGAUUUCUGCCCAGUGCUCUGAAUGUCAAAGU
GAAGAAAUUCAAUGAAGCGCGGGUAAACGGCGGGAGUAACUAUGACUCUCUUAAGGUAGC
CAAAUGCCUCGUCAUCUAAUUAGUGACGCGCAUGAAXGGAUGAACGAGAUUCCCACUGUC
CCUACCUACUAUCCAGCGAAACCACAGCCAAGGGAACGGGCUUGGCGGAAUCAGCGGGGA
AAGAAGACCCUGUUGAGCUUGACUCUAGUCUGGCACGGUGAAGAGACAUGAGAGGUGUAG
AAUAAGUGGGAGGCCCCCGGCGCCCCCCCGGUGUCCCCGCGAGGGGCCCGGGGCGGGGUC
CGCCGGCCCUGCGGGCCGCCGGUGAAAUACCACUACUCUGAUCGUUUUUUCACUGACCCG
GUGAGGCGGGGGGGCGAGCCCCGAGGGGCUCUCGCUUCUGGCGCCAAGCGCCCGGCCGCG
CGCCGGCCGGGCGCGACCCGCUCCGGGGACAGUGCCAGGUGGGGAGUUUGACUGGGGCGG
UACACCUGUCAAACGGUAACGCAGGUGUCCUAAGGCGAGCUCAGGGAGGACAGAAACCUC
CCGUGGAGCAGAAGGGCAAAAGCUCGCUUGAUCUUGAUUUUCAGUACGAAUACAGACCGU
GAAAGCGGGGCCUUACGAUCCUUCUGACCUUUUGGGUUUUAAGCAGGAGGUGUCAGAAAA
GUUACCACAGGGAUAACUGGCUUGUGGCGGCCAAGCGUUCAUAGCGACGUCGCUUUUUGA
UCCUUCGAUGUCGGCUCUUCCUAUCAUUGUGAAGCAGAAUUCACCAAGCGUUGGAUUGUU
CACCCACUAAUAGGGAACGUGAGCUGGGUUUAGACCGUCGUGAGACAGGUUAGUUUUACC
CUACUGAUGAUGUGUUGUUGCCAUGGUAAUCCUGCUCAGUACGAGAGGAACCGCAGGUUC
AGACAUUUGGUGUAUGUGCUUGGCUGAGGAGCCAAUGGGGCGAAGCUACCAUCUGUGGGA
UUAUGACUGAACGCCUCUAAGUCAGAAUCCCGCCCAGGCGGAACGAUACGGCAGCGCCGC
GGAGCCUCGGUUGGCCUCGGAUAGCCGGUCCCCCGCCUGUCCCCGCCGGCGGGCCGCCCC
CCCCCUCCACGCGCCCCGCGCGCGCGGGAGGGCGCGUGCCCCGCCGCGCGCCGGGACCGG
GGUCCGGUGCGGAGUGCCCUUCGUCCUGGGAAACGGGGCGCGGCCGGAAAGGCGGCCGCC
CCCUCGCCCGUCACGCACCGCACGUUCGUGGGGAACCUGGCGCUAAACCAUUCGUAGACG
ACCUGCUUCUGGGUCGGGGU
Sequence of entity 5 (B7), FASTA
>28LN_5 5S rRNA (chains B7)
GUCUACGGCCAUACCACCCUGAACGCGCCCGAUCUCGUCUGAUCUCGGAAGCUAAGCAGG
GUCGGGCCUGGUUAGUACUUGGAUGGGAGACCGCCUGGGAAUACCGGGUGCUGUAGGCUU
Sequence of entity 6 (B8), FASTA
>28LN_6 5.8S rRNA (chains B8)
CGACUCUUAGCGGUGGAUCACUCGGCUCGUGCGUCGAUGAAGAACGCAGCUAGCUGCGAG
AAUUAAUGUGAAUUGCAGGACACAUUGAUCAUCGACACUUCGAACGCACUUGCGGCCCCG
GGUUCCUCCCGGGGCUACGCCUGUCUGAGCGUCGCUU
Sequence of entity 7 (BA), FASTA
>28LN_7 Large ribosomal subunit protein uL2 (chains BA)
MGRVIRGQRKGAGSVFRAHVKHRKGAARLRAVDFAERHGYIKGIVKDIIHDPGRGAPLAK
VVFRDPYRFKKRTELFIAAEGIHTGQFVYCGKKAQLNIGNVLPVGTMPEGTIVCCLEEKP
GDRGKLARASGNYATVISHNPETKKTRVKLPSGSKKVISSANRAVVGVVAGGGRIDKPIL
KAGRAYHKYKAKRNCWPRVRGVAMNPVEHPFGGGNXQHIGKPSTIRRDAPAGRKVGLIAA
RRTGRLRGTKTVQEKEN
Sequence of entity 8 (BB), FASTA
>28LN_8 Large ribosomal subunit protein uL3 (chains BB)
MSHRKFSAPRHGSLGFLPRKRSSRHRGKVKSFPKDDPSKPVHLTAFLGYKAGMTHIVREV
DRPGSKVNKKEVVEAVTIVETPPMVVVGIVGYVETPRGLRTFKTVFAEHISDECKRRFYK
NWHKSKKKAFTKYCKKWQDEDGKKQLEKDFSSMKKYCQVIRVIAHTQMRLLPLRQKKAHL
MEIQVNGGTVAEKLDWARERLEQQVPVNQVFGQDEMIDVIGVTKGKGYKGVTSRWHTKKL
PRKTHRGLRKVACIGAWHPARVAFSVARAGQKGYHHRTEINKKIYKIGQGYLIKDGKLIK
NNASTDYDLSDKSINPLGGFVHYGEVTNDFVMLKGCVVGTKKRVLTLRKSLLVQTKRRAL
EKIDLKFIDTTSKFGHGRFQTMEEKKAFMGPLKKDRIAKEEGA
Sequence of entity 9 (BC), FASTA
>28LN_9 60S ribosomal protein L4 (chains BC)
MACARPLISVYSEKGESSGKNVTLPAVFKAPIRPDIVNFVHTNLRKNNRQPYAVSELAGH
QTSAESWGTGRAVARIPRVRGGGTHRSGQGAFGNMCRGGRMFAPTKTWRRWHRRVNTTQK
RYAICSALAASALPALVMSKGHRIEEVPELPLVVEDKVEGYKKTKEAVLLLKKLKAWNDI
KKVYASQRMRAGKGKMRNRRRIQRRGPCIIYNEDNGIIKAFRNIPGITLLNVSKLNILKL
APGGHVGRFCIWTESAFRKLDELYGTWRKAASLKSNYNLPMHKMINTDLSRILKSPEIQR
ALRAPRKKIHRRVLKKNPLKNLRIMLKLNPYAKTMRRNTILRQARNHKLRVDKAAAAAAA
LQAKSDEKAAVAGKKPVVGKKGKKAAVGVKKQKKPLVGKKAAATKKPAPEKKPAEKKPTT
EEKKPAA
Sequence of entity 10 (BD), FASTA
>28LN_10 60S ribosomal protein L5 (chains BD)
MGFVKVVKNKAYFKRYQVKFRRRREGKTDYYARKRLVIQDKNKYNTPKYRMIVRVTNRDI
ICQIAYARIEGDMIVCAAYAHELPKYGVKVGLTNYAAAYCTGLLLARRLLNRFGMDKIYE
GQVEVTGDEYNVESIDGQPGAFTCYLDAGLARTTTGNKVFGALKGAVDGGLSIPHSTKRF
PGYDSESKEFNAEVHRKHIMGQNVADYMRYLMEEDEDAYKKQFSQYIKNSVTPDMMEEMY
KKAHAAIRENPVYEKKPKKEVKKKRWNRPKMSLAQKKDRVAQKKASFLRAQERAAES
Sequence of entity 11 (BE), FASTA
>28LN_11 Large ribosomal subunit protein eL6 (chains BE)
MAGEKVEKPDTKEKKPEAKKVDAGGKVKKGNLKAKKPKKGKPHCSRNPVLVRGIGRYSRS
AMYSRKAMYKRKYSAAKSKVEKKKKEKVLATVTKPVGGDKNGGTRVVKLRKMPRYYPTED
VPRKLLSHGKKPFSQHVRKLRASITPGTILIILTGRHRGKRVVFLKQLASGLLLVTGPLV
LNRVPLRRTHQKFVIATSTKIDISNVKIPKHLTDAYFKKKKLRKPRHQEGEIFDTEKEKY
EITEQRKIDQKAVDSQILPKIKAIPQLQGYLRSVFALTNGIYPHKLVF
Sequence of entity 12 (BF), FASTA
>28LN_12 Large ribosomal subunit protein uL30 (chains BF)
MEGVEEKKKEVPAVPETLKKKRRNFAELKIKRLRKKFAQKMLRKARRKLIYEKAKHYHKE
YRQMYRTEIRMARMARKAGNFYVPAEPKLAFVIRIRGINGVSPKVRKVLQLLRLRQIFNG
TFVKLNKASINMLRIVEPYIAWGYPNLKSVNELIYKRGYGKINKKRIALTDNALIARSLG
KYGIICMEDLIHEIYTVGKRFKEANNFLWPFKLSSPRGGMKKKTTHFVEGGDAGNREDQI
NRLIRRMN

Ligands and cofactors

IDNameFormulaCopies
MGMagnesium ionMg432
ZNZinc ionZn8

Primary citation

NAC promotes co-translational protein folding at the ribosomal tunnel exit. Santos, J., Gunnigmann, M., Gora, R.J. et al. Mol Cell (2026) 86:1311-1326.e11. DOI 10.1016/j.molcel.2026.02.022 · PubMed

Other PDB entries of the same protein (UniProt P17861 (AlphaFold model), which also has an AlphaFold model), best resolution first:

Browse structure collections

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