9H52: GTPase Era, mitochondrial

Assembly intermediate of human mitochondrial ribosome small subunit in complex with NOA1, ERAL1, METTL17, MCAT and TFB1M (state N1). Determined by electron microscopy at 3.9 Å resolution. Released 8 Jul 2026.

Method
Electron microscopy
Resolution
3.9 Å
Organism
Homo sapiens
Chains
30
Atoms
67,748
Mol. weight
1296.84 kDa
Ligands
ZN, FES, MG, ATP
Released
8 Jul 2026

Explore 9H52 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9H52 contains 366 α-helices and 232 β-strands across 28 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 0: 15 helices, 15 β-strands

ElementResiduesLengthSheet
α-helix10-2516
α-helix27-293
α-helix30-334
β-strand3718
β-strand4218
α-helix481
β-strand4918
α-helix50-512
α-helix52-587
β-strand6219
α-helix64-685
α-helix72-754
β-strand79-81310
α-helix83-886
β-strand94-97410
β-strand101-102211
β-strand112-113211
β-strand115-120610
β-strand123-124210
β-strand129-130210
α-helix1311
β-strand13919
β-strand140-141210
α-helix147-1504
β-strand163-165312
α-helix166-1672
α-helix168-18013
α-helix187-1893
β-strand190-192312
α-helix213-2153
Chain 1: 17 helices, 5 β-strands
ElementResiduesLengthSheet
α-helix66-694
β-strand86159
α-helix89-913
α-helix96-983
α-helix103-1064
α-helix116-12510
α-helix126-1294
α-helix140-1467
β-strand150-152370
α-helix166-1694
β-strand171-176670
α-helix177-1793
α-helix184-19411
α-helix195-1973
β-strand198-199270
β-strand204-209670
α-helix215-23420
α-helix240-2423
α-helix257-27317
α-helix283-2853
α-helix287-29812
α-helix306-31914
Chain 4: 41 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix59-635
α-helix69-779
α-helix100-12324
α-helix125-1273
α-helix138-1403
α-helix154-1629
α-helix166-17813
α-helix181-1844
α-helix185-19814
α-helix236-2394
α-helix243-2508
α-helix256-26813
α-helix272-28413
α-helix291-30313
α-helix309-32517
α-helix328-3303
α-helix332-34211
α-helix343-3497
α-helix350-36314
α-helix365-3673
α-helix369-37911
α-helix392-4009
α-helix412-4143
α-helix415-42612
α-helix429-43911
α-helix444-4485
α-helix451-46818
α-helix471-48111
α-helix487-4893
α-helix490-50314
α-helix506-5083
α-helix509-51810
α-helix525-53612
α-helix542-56019
α-helix573-58513
α-helix589-60214
α-helix605-6073
α-helix608-62114
α-helix624-63613
α-helix642-65211
α-helix657-66610
Chain 5: 19 helices, 7 β-strands
ElementResiduesLengthSheet
α-helix13-175
α-helix18-258
α-helix30-334
α-helix41-5111
β-strand58-62525
α-helix68-758
β-strand80-85625
α-helix91-10010
β-strand105-109525
α-helix123-1253
β-strand136-140525
α-helix144-16017
α-helix163-1675
β-strand171-177725
α-helix178-1847
α-helix195-2017
β-strand204-212925
α-helix214-2163
β-strand225-232825
α-helix242-25312
α-helix260-2656
α-helix270-28415
α-helix292-2943
α-helix297-31216
α-helix315-3195
α-helix322-3276
Chain 7: 20 helices, 16 β-strands
ElementResiduesLengthSheet
β-strand36-37213
α-helix43-464
α-helix49-513
α-helix57-604
β-strand61-62214
α-helix65-7511
α-helix81-9616
α-helix97-1026
α-helix103-11513
α-helix134-1418
α-helix152-1587
α-helix159-1635
α-helix164-18017
β-strand189-192414
α-helix198-20710
β-strand214-217414
α-helix221-23111
β-strand245-248414
β-strand259-265714
α-helix268-2703
α-helix274-28714
β-strand288-2971014
α-helix300-31415
β-strand327-331514
α-helix338-3392
β-strand349-354614
α-helix355-3573
β-strand365-3761214
β-strand387-388215
β-strand393-395315
β-strand398-404715
β-strand410-416715
α-helix417-4204
α-helix422-4309
β-strand436115
β-strand439-440213
Chain 8: 19 helices, 11 β-strands
ElementResiduesLengthSheet
α-helix57-582
β-strand63-6756
α-helix87-9812
α-helix102-1076
α-helix117-13822
α-helix140-1445
β-strand146-15166
α-helix155-1628
α-helix168-18821
β-strand192-19877
α-helix204-21815
β-strand225-23287
β-strand235-24067
α-helix242-2509
α-helix252-2554
β-strand257-26377
α-helix272-2776
α-helix278-28710
β-strand296-29836
α-helix2991
β-strand30516
α-helix309-3113
α-helix312-3187
α-helix319-3213
β-strand324-32527
α-helix326-3349
β-strand344-34746
α-helix353-3619
α-helix363-3664
β-strand369-37136
Chain 9: 24 helices, 30 β-strands
ElementResiduesLengthSheet
α-helix75-773
α-helix88-11427
α-helix180-19314
β-strand195-196216
α-helix198-2003
α-helix201-2055
α-helix206-2094
β-strand215-222817
α-helix231-2333
α-helix234-2385
β-strand242-248717
α-helix260-27415
β-strand278-279217
β-strand309-315717
β-strand317118
β-strand322118
α-helix324-3318
β-strand340-344517
β-strand346116
α-helix351-3599
β-strand364119
α-helix371-3744
β-strand376-379420
α-helix3801
β-strand386-389420
β-strand391121
β-strand392119
α-helix393-3953
α-helix397-4004
β-strand493121
β-strand494-496317
α-helix497-4982
β-strand499-500216
α-helix5011
α-helix506-5094
α-helix512-5187
β-strand527-531522
β-strand535-539523
β-strand543-548623
β-strand554-559622
β-strand566-570523
α-helix571-58111
α-helix593-5997
α-helix601-6022
β-strand603-609723
α-helix610-6134
β-strand621-624422
β-strand630-634522
β-strand642-648723
β-strand654-656322
β-strand670124
β-strand676122
β-strand677124
α-helix681-6844
Chain a: 15 helices, 16 β-strands
ElementResiduesLengthSheet
α-helix94-1018
β-strand112-11981
α-helix126-1349
β-strand152-15871
β-strand161-16771
α-helix168-1703
α-helix189-1946
β-strand199-20571
α-helix217-2259
β-strand231-23661
α-helix238-2403
α-helix246-25510
β-strand258-25922
β-strand262-26322
α-helix2641
β-strand26613
β-strand302-30761
β-strand30914
β-strand31414
α-helix316-32611
α-helix3271
β-strand328-32921
β-strand34011
α-helix344-35916
α-helix364-3674
β-strand369-37795
β-strand383-39085
α-helix394-4018
α-helix403-4053
α-helix406-42318
β-strand427-43375

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

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
GTPase Era, mitochondrialaprotein437Homo sapiensO75616 (AlphaFold model)
Small ribosomal subunit protein mS340protein215Homo sapiensP82930 (AlphaFold model)
Nitric oxide-associated protein 19protein698Homo sapiensQ8NC60 (AlphaFold model)
28S ribosomal protein S5, mitochondrialDprotein430Homo sapiensP82675 (AlphaFold model)
28S ribosomal protein S6, mitochondrialEprotein125Homo sapiensP82932
28S ribosomal protein S12, mitochondrialJprotein138Homo sapiensO15235
28S ribosomal protein S15, mitochondrialLprotein257Homo sapiensP82914
28S ribosomal protein S16, mitochondrialMprotein137Homo sapiensQ9Y3D3
28S ribosomal protein S17, mitochondrialNprotein130Homo sapiensQ9Y2R5
28S ribosomal protein S18b, mitochondrialOprotein258Homo sapiensQ9Y676
28S ribosomal protein S18c, mitochondrialPprotein142Homo sapiensQ9Y3D5
MRPS21 isoform 1Qprotein87Homo sapiensA0A2J8VEN6

18 more molecules are not listed.

Sequence of entity 1 (a), FASTA
>9H52_1 GTPase Era, mitochondrial (chains a)
MAAPSWRGARLVQSVLRVWQVGPHVARERVIPFSSLLGFQRRCVSCVAGSAFSGPRLASA
SRSNGQGSALDHFLGFSQPDSSVTPCVPAVSMNRDEQDVLLVHHPDMPENSRVLRVVLLG
APNAGKSTLSNQLLGRKVFPVSRKVHTTRCQALGVITEKETQVILLDTPGIISPGKQKRH
HLELSLLEDPWKSMESADLVVVLVDVSDKWTRNQLSPQLLRCLTKYSQIPSVLVMNKVDC
LKQKSVLLELTAALTEGVVNGKKLKMRQAFHSHPGTHCPSPAVKDPNTQSVGNPQRIGWP
HFKEIFMLSALSQEDVKTLKQYLLTQAQPGPWEYHSAVLTSQTPEEICANIIREKLLEHL
PQEVPYNVQQKTAVWEEGPGGELVIQQKLLVPKESYVKLLIGPKGHVISQIAQEAGHDLM
DIFLCDVDIRLSVKLLK
Sequence of entity 2 (0), FASTA
>9H52_2 Small ribosomal subunit protein mS34 (chains 0)
KKVRPRLIAELARRVRALREQLNRPRDSQLYAVDYETLTRPFSGRRLPVRAWADVRRESR
LLQLLGRLPLFGLGRLVTRKSWLWQHDEPCYWRLTRVRPDYTAQNLDHGKAWGILTFKGK
TESEAREIEHVMYHDWRLVPKHEEEAFTAFTPAPEDSLASVPYPPLLRAMIIAERQKNGD
TSTEEPMLNVQRIRMEPWDYPAKQEDKGRAKGTPV
Sequence of entity 3 (9), FASTA
>9H52_3 Nitric oxide-associated protein 1 (chains 9)
MLPARLPFRLLSLFLRGSAPTAARHGLREPLLERRCAAASSFQHSSSLGRELPYDPVDTE
GFGEGGDMQERFLFPEYILDPEPQPTREKQLQELQQQQEEEERQRQQRREERRQQNLRAR
SREHPVVGHPDPALPPSGVNCSGCGAELHCQDAGVPGYLPREKFLRTAEADGGLARTVCQ
RCWLLSHHRRALRLQVSREQYLELVSAALRRPGPSLVLYMVDLLDLPDALLPDLPALVGP
KQLIVLGNKVDLLPQDAPGYRQRLRERLWEDCARAGLLLAPGHQGPQRPVKDEPQDGENP
NPPNWSRTVVRDVRLISAKTGYGVEELISALQRSWRYRGDVYLVGATNAGKSTLFNTLLE
SDYCTAKGSEAIDRATISPWPGTTLNLLKFPICNPTPYRMFKRHQRLKKDSTQAEEDLSE
QEQNQLNVLKKHGYVVGRVGRTFLYSEEQKDNIPFEFDADSLAFDMENDPVMGTHKSTKQ
VELTAQDVKDAHWFYDTPGITKENCILNLLTEKEVNIVLPTQSIVPRTFVLKPGMVLFLG
AIGRIDFLQGNQSAWFTVVASNILPVHITSLDRADALYQKHAGHTLLQIPMGGKERMAGF
PPLVAEDIMLKEGLGASEAVADIKFSSAGWVSVTPNFKDRLHLRGYTPEGTVLTVRPPLL
PYIVNIKGQRIKKSVAYKTKKPPSLMYNVRKKKGKINV
Sequence of entity 4 (D), FASTA
>9H52_4 28S ribosomal protein S5, mitochondrial (chains D)
MATAVRAVGCLPVLCSGTAGHLLGRQCSLNTLPAASILAWKSVLGNGHLSSLGTRDTHPY
ASLSRALQTQCCISSPSHLMSQQYRPYSFFTKLTADELWKGALAETGAGAKKGRGKRTKK
KKRKDLNRGQIIGEGRYGFLWPGLNVPLMKNGAVQTIAQRSKEEQEKVEADMIQQREEWD
RKKKMKVKRERGWSGNSWGGISLGPPDPGPCGETYEDFDTRILEVRNVFTMTAKEGRKKS
IRVLVAVGNGKGAAGFSIGKATDRMDAFRKAKNRAVHHLHYIERYEDHTIFHDISLRFKR
THIKMKKQPKGYGLRCHRAIITICRLIGIKDMYAKVSGSINMLSLTQGLFRGLSRQETHQ
QLADKKGLHVVEIREECGPLPIVVASPRGPLRKDPEPEDEVPDVKLDWEDVKTAQGMKRS
VWSNLKRAAT
Sequence of entity 5 (E), FASTA
>9H52_5 28S ribosomal protein S6, mitochondrial (chains E)
MPRYELALILKAMQRPETAATLKRTIEALMDRGAIVRDLENLGERALPYRISAHSQQHNR
GGYFLVDFYAPTAAVESMVEHLSRDIDVIRGNIVKHPLTQELKECEGIVPVPLAEKLYST
KKRKK
Sequence of entity 6 (J), FASTA
>9H52_6 28S ribosomal protein S12, mitochondrial (chains J)
MSWSGLLHGLNTSLTCGPALVPRLWATCSMATLNQMHRLGPPKRPPRKLGPTEGRPQLKG
VVLCTFTRKPKKPNSANRKCCRVRLSTGREAVCFIPGEGHTLQEHQIVLVEGGRTQDLPG
VKLTVVRGKYDCGHVQKK
Sequence of entity 7 (L), FASTA
>9H52_7 28S ribosomal protein S15, mitochondrial (chains L)
MLRVAWRTLSLIRTRAVTQVLVPGLPGGGSAKFPFNQWGLQPRSLLLQAARGYVVRKPAQ
SRLDDDPPPSTLLKDYQNVPGIEKVDDVVKRLLSLEMANKKEMLKIKQEQFMKKIVANPE
DTRSLEARIIALSVKIRSYEEHLEKHRKDKAHKRYLLMSIDQRKKMLKNLRNTNYDVFEK
ICWGLGIEYTFPPLYYRRAHRRFVTKKALCIRVFQETQKLKKRRRALKAAAAAQKQAKRR
NPDSPAKAIPKTLKDSQ
Sequence of entity 8 (M), FASTA
>9H52_8 28S ribosomal protein S16, mitochondrial (chains M)
MVHLTTLLCKAYRGGHLTIRLALGGCTNRPFYRIVAAHNKCPRDGRFVEQLGSYDPLPNS
HGEKLVALNLDRIRHWIGCGAHLSKPMEKLLGLAGFFPLHPMMITNAERLRRKRAREVLL
ASQKTDAEATDTEATET
Sequence of entity 9 (N), FASTA
>9H52_9 28S ribosomal protein S17, mitochondrial (chains N)
MSVVRSSVHARWIVGKVIGTKMQKTAKVRVTRLVLDPYLLKYFNKRKTYFAHDALQQCTV
GDIVLLRALPVPRAKHVKHELAEIVFKVGKVIDPVTGKPCAGTTYLESPLSSETTQLSKN
LEELNISSAQ
Sequence of entity 10 (O), FASTA
>9H52_10 28S ribosomal protein S18b, mitochondrial (chains O)
MAASVLNTVLRRLPMLSLFRGSHRVQVPLQTLCTKAPSEEDSLSSVPISPYKDEPWKYLE
SEEYQERYGSRPVWADYRRNHKGGVPPQRTRKTCIRRNKVVGNPCPICRDHKLHVDFRNV
KLLEQFVCAHTGIIFYAPYTGVCVKQHKRLTQAIQKARDHGLLIYHIPQVEPRDLDFSTS
HGAVSATPPAPTLVSGDPWYPWYNWKQPPERELSRLRRLYQGHLQEESGPPPESMPKMPP
RTPAEASSTGQTGPQSAL
Sequence of entity 11 (P), FASTA
>9H52_11 28S ribosomal protein S18c, mitochondrial (chains P)
MAAVVAVCGGLGRKKLTHLVTAAVSLTHPGTHTVLWRRGCSQQVSSNEDLPISMENPYKE
PLKKCILCGKHVDYKNVQLLSQFVSPFTGCIYGRHITGLCGKKQKEITKAIKRAQIMGFM
PVTYKDPAYLKDPKVCNIRYRE
Sequence of entity 12 (Q), FASTA
>9H52_12 MRPS21 isoform 1 (chains Q)
XAKHLKFIARTVMVQEGNVESAYRTLNRILTMDGLIEDIKHRRYYEKPCRRRQRESYERC
RRIYNMEMARKINFLMRKNRADPWQGC

Ligands and cofactors

IDNameFormulaCopies
ZNZinc ionZn1
FESFE2/S2 (inorganic) clusterFe2 S22
MGMagnesium ionMg2
ATPAdenosine-5'-triphosphateC10 H16 N5 O13 P31
SF4Iron/sulfur clusterFe4 S41
GDPGuanosine-5'-diphosphateC10 H15 N5 O11 P22

Primary citation

Pseudouridine synthase PUS1 and initiation factor mtIF2 are human mitoribosomal small subunit assembly factors. Singh, V., Shiriaev, D., Bilalli, L. et al. Nat Commun (2026) 17. DOI 10.1038/s41467-026-74700-x · PubMed

Other PDB entries of the same protein (UniProt O75616 (AlphaFold model), which also has an AlphaFold model), best resolution first:

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