9PR4: 39S ribosomal protein L32, mitochondrial

In Situ Structure of the Human Mitochondrial Large Subunit 39S in Complex with TACO1. Determined by electron microscopy at 2.77 Å resolution. Released 18 Feb 2026.

Method: Electron microscopy
Resolution: 2.77 Å
Organism: Homo sapiens
Chains: 63
Atoms: 111,602
Mol. weight: 2017.65 kDa
Ligands: ZN, MG, FES, SPD
Released: 18 Feb 2026

Explore 9PR4 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9PR4 contains 520 α-helices and 433 β-strands across 60 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: 5 helices, 4 β-strands

Element	Residues	Length	Sheet
α-helix	84-86	3
α-helix	87-94	8
α-helix	101-104	4
β-strand	107-109	3	1
β-strand	116-118	3	1
α-helix	124-145	22
β-strand	156-159	4	2
β-strand	173-176	4	2
α-helix	180-182	3

Chain 1: 0 helices, 4 β-strands

Element	Residues	Length	Sheet
β-strand	15-22	8	6
β-strand	29-35	7	6
β-strand	42-47	6	6
β-strand	52-64	13	6

Chain 2: 3 helices, 0 β-strands

Element	Residues	Length
α-helix	57-64	8
α-helix	66-71	6
α-helix	73-85	13

Chain 3: 6 helices, 4 β-strands

Element	Residues	Length	Sheet
α-helix	108-111	4
β-strand	115-116	2	15
β-strand	122-125	4	15
α-helix	133-135	3
α-helix	138-144	7
β-strand	147-149	3	15
α-helix	152-160	9
α-helix	164-167	4
β-strand	172	1	16
α-helix	178-181	4

Chain 4: 1 helix, 4 β-strands

Element	Residues	Length	Sheet
β-strand	67-69	3	23
β-strand	80-84	5	23
β-strand	87-91	5	23
α-helix	96-98	3
β-strand	100-102	3	23

Chain 5: 24 helices, 20 β-strands

Element	Residues	Length	Sheet
α-helix	31-32	2
α-helix	40-47	8
α-helix	48-51	4
β-strand	59	1	28
β-strand	71	1	28
α-helix	72-73	2
α-helix	75-77	3
α-helix	85-87	3
α-helix	90-92	3
α-helix	93-95	3
β-strand	99-107	9	29
α-helix	117-124	8
β-strand	126-130	5	30
α-helix	134-141	8
α-helix	143-145	3
α-helix	147	1
α-helix	150-159	10
α-helix	160-164	5
α-helix	173-189	17
α-helix	190-192	3
α-helix	196-200	5
β-strand	201-205	5	30
β-strand	209-215	7	29
β-strand	218-224	7	29
β-strand	228-232	5	30
α-helix	235-238	4
α-helix	242-246	5
α-helix	247-249	3
α-helix	253-255	3
β-strand	259	1	31
β-strand	266-270	5	29
β-strand	276	1	29
β-strand	279	1	32
β-strand	287-294	8	30
α-helix	304-305	2
α-helix	306-329	24
β-strand	335-347	13	30
β-strand	351-361	11	30
β-strand	372-381	10	30
β-strand	385-387	3	33
β-strand	391-393	3	34
β-strand	396-399	4	34
β-strand	402-404	3	33
α-helix	408-419	12

Chain 6: 22 helices, 16 β-strands

Element	Residues	Length	Sheet
α-helix	34-35	2
α-helix	49	1
β-strand	50	1	43
α-helix	54-66	13
α-helix	74-78	5
α-helix	86-88	3
β-strand	89	1	44
α-helix	92-96	5
α-helix	99-115	17
α-helix	117-124	8
α-helix	132-142	11
α-helix	144-154	11
α-helix	157-161	5
β-strand	166	1	44
β-strand	173-179	7	45
β-strand	183-187	5	45
β-strand	191-192	2	46
β-strand	203-206	4	45
β-strand	213-220	8	46
β-strand	231-241	11	46
α-helix	245-247	3
β-strand	249-252	4	46
α-helix	259-260	2
β-strand	261	1	47
β-strand	266-275	10	46
α-helix	282-284	3
α-helix	293-296	4
β-strand	297-298	2	46
α-helix	300-305	6
β-strand	311-321	11	46
α-helix	325-329	5
α-helix	330-334	5
α-helix	338-339	2
β-strand	340-345	6	47
α-helix	346-350	5
α-helix	352-354	3
β-strand	359	1	15
α-helix	363-369	7

Chain 7: 14 helices, 17 β-strands

Element	Residues	Length	Sheet
α-helix	37-57	21
β-strand	64-69	6	50
β-strand	78-82	5	50
α-helix	88-95	8
α-helix	97-102	6
β-strand	105-108	4	51
β-strand	111-113	3	51
α-helix	118-119	2
β-strand	123-125	3	50
β-strand	127-128	2	51
α-helix	136-155	20
β-strand	165-168	4	52
α-helix	174-176	3
β-strand	179-184	6	52
α-helix	187-189	3
α-helix	196-212	17
β-strand	215	1	53
β-strand	217-222	6	54
α-helix	223-229	7
α-helix	234-246	13
β-strand	251-257	7	54
β-strand	260-263	4	54
β-strand	271	1	53
α-helix	272-274	3
β-strand	275-279	5	55
β-strand	283-285	3	52
β-strand	294-299	6	52
β-strand	300-303	4	55
α-helix	304-306	3
α-helix	307-309	3
α-helix	310-320	11

52 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
39S ribosomal protein L32, mitochondrial	0	protein	188	Homo sapiens	Q9BYC8 (AlphaFold model)
39S ribosomal protein L33, mitochondrial	1	protein	65	Homo sapiens	O75394 (AlphaFold model)
39S ribosomal protein L34, mitochondrial	2	protein	92	Homo sapiens	Q9BQ48 (AlphaFold model)
39S ribosomal protein L35, mitochondrial	3	protein	188	Homo sapiens	Q9NZE8 (AlphaFold model)
39S ribosomal protein L36, mitochondrial	4	protein	103	Homo sapiens	Q9P0J6
39S ribosomal protein L37, mitochondrial	5	protein	423	Homo sapiens	Q9BZE1
39S ribosomal protein L38, mitochondrial	6	protein	380	Homo sapiens	Q96DV4
39S ribosomal protein L39, mitochondrial	7	protein	338	Homo sapiens	Q9NYK5
39S ribosomal protein L40, mitochondrial	8	protein	206	Homo sapiens	Q9NQ50
39S ribosomal protein L41, mitochondrial	9	protein	137	Homo sapiens	Q8IXM3
Translational activator of cytochrome c oxidase 1	C	protein	297	Homo sapiens	Q9BSH4
39S ribosomal protein L2, mitochondrial	D	protein	305	Homo sapiens	Q5T653

45 more molecules are not listed.

Sequence of entity 1 (0), FASTA

>9PR4_1 39S ribosomal protein L32, mitochondrial (chains 0)
MALAMLVLVVSPWSAARGVLRNYWERLLRKLPQSRPGFPSPPWGPALAVQGPAMFTEPAN
DTSGSKENSSLLDSIFWMAAPKNRRTIEVNRCRRRNPQKLIKVKNNIDVCPECGHLKQKH
VLCAYCYEKVCKETAEIRRQIGKQEGGPFKAPTIETVVLYTGETPSEQDQGKRIIERDRK
RPSWFTQN

Sequence of entity 2 (1), FASTA

>9PR4_2 39S ribosomal protein L33, mitochondrial (chains 1)
MFLSAVFFAKSKSKNILVRMVSEAGTGFCFNTKRNRLREKLTLLHYDPVVKQRVLFVEKK
KIRSL

Sequence of entity 3 (2), FASTA

>9PR4_3 39S ribosomal protein L34, mitochondrial (chains 2)
MAVLAGSLLGPTSRSAALLGGRWLQPRAWLGFPDAWGLPTPQQARGKARGNEYQPSNIKR
KNKHGWVRRLSTPAGVQVILRRMLKGRKSLSH

Sequence of entity 4 (3), FASTA

>9PR4_4 39S ribosomal protein L35, mitochondrial (chains 3)
MAASAFAGAVRAASGILRPLNILASSTYRNCVKNASLISALSTGRFSHIQTPVVSSTPRL
TTSERNLTCGHTSVILNRMAPVLPSVLKLPVRSLTYFSARKGKRKTVKAVIDRFLRLHCG
LWVRRKAGYKKKLWKKTPARKKRLREFVFCNKTQSKLLDKMTTSFWKRRNWYVDDPYQKY
HDRTNLKV

Sequence of entity 5 (4), FASTA

>9PR4_5 39S ribosomal protein L36, mitochondrial (chains 4)
MANLFIRKMVNPLLYLSRHTVKPRALSTFLFGSIRGAAPVAVEPGAAVRSLLSPGLLPHL
LPALGFKNKTVLKKRCKDCYLVKRRGRWYVYCKTHPRHKQRQM

Sequence of entity 6 (5), FASTA

>9PR4_6 39S ribosomal protein L37, mitochondrial (chains 5)
MALASGPARRALAGSGQLGLGGFGAPRRGAYEWGVRSTRKSEPPPLDRVYEIPGLEPITF
AGKMHFVPWLARPIFPPWDRGYKDPRFYRSPPLHEHPLYKDQACYIFHHRCRLLEGVKQA
LWLTKTKLIEGLPEKVLSLVDDPRNHIENQDECVLNVISHARLWQTTEEIPKRETYCPVI
VDNLIQLCKSQILKHPSLARRICVQNSTFSATWNRESLLLQVRGSGGARLSTKDPLPTIA
SREEIEATKNHVLETFYPISPIIDLHECNIYDVKNDTGFQEGYPYPYPHTLYLLDKANLR
PHRLQPDQLRAKMILFAFGSALAQARLLYGNDAKVLEQPVVVQSVGTDGRVFHFLVFQLN
TTDLDCNEGVKNLAWVDSDQLLYQHFWCLPVIKKRVVVEPVGPVGFKPETFRKFLALYLH
GAA

Sequence of entity 7 (6), FASTA

>9PR4_7 39S ribosomal protein L38, mitochondrial (chains 6)
MAAPWWRAALCECRRWRGFSTSAVLGRRTPPLGPMPNSDIDLSNLERLEKYRSFDRYRRR
AEQEAQAPHWWRTYREYFGEKTDPKEKIDIGLPPPKVSRTQQLLERKQAIQELRANVEEE
RAARLRTASVPLDAVRAEWERTCGPYHKQRLAEYYGLYRDLFHGATFVPRVPLHVAYAVG
EDDLMPVYCGNEVTPTEAAQAPEVTYEAEEGSLWTLLLTSLDGHLLEPDAEYLHWLLTNI
PGNRVAEGQVTCPYLPPFPARGSGIHRLAFLLFKQDQPIDFSEDARPSPCYQLAQRTFRT
FDFYKKHQETMTPAGLSFFQCRWDDSVTYIFHQLLDMREPVFEFVRPPPYHPKQKRFPHR
QPLRYLDRYRDSHEPTYGIY

Sequence of entity 8 (7), FASTA

>9PR4_8 39S ribosomal protein L39, mitochondrial (chains 7)
MEALAMGSRALRLWLVAPGGGIKWRFIATSSASQLSPTELTEMRNDLFNKEKARQLSLTP
RTEKIEVKHVGKTDPGTVFVMNKNISTPYSCAMHLSEWYCRKSILALVDGQPWDMYKPLT
KSCEIKFLTFKDCDPGEVNKAYWRSCAMMMGCVIERAFKDEYMVNLVRAPEVPVISGAFC
YDVVLDSKLDEWMPTKENLRSFTKDAHALIYKDLPFETLEVEAKVALEIFQHSKYKVDFI
EEKASQNPERIVKLHRIGDFIDVSEGPLIPRTSICFQYEVSAVHNLQPTQPSLIRRFQGV
SLPVHLRAHFTIWDKLLERSRKMVTEDQSKATEECTST

Sequence of entity 9 (8), FASTA

>9PR4_9 39S ribosomal protein L40, mitochondrial (chains 8)
MTASVLRSISLALRPTSGLLGTWQTQLRETHQRASLLSFWELIPMRSEPLRKKKKVDPKK
DQEAKERLKRKIRKLEKATQELIPIEDFITPLKFLDKARERPQVELTFEETERRALLLKK
WSLYKQQERKMERDTIRAMLEAQQEALEELQLESPKLHAEAIKRDPNLFPFEKEGPHYTP
PIPNYQPPEGRYNDITKVYTQVEFKR

Sequence of entity 10 (9), FASTA

>9PR4_10 39S ribosomal protein L41, mitochondrial (chains 9)
MGVLAAAARCLVRGADRMSKWTSKRGPRSFRGRKGRGAKGIGFLTSGWRFVQIKEMVPEF
VVPDLTGFKLKPYVSYLAPESEETPLTAAQLFSEAVAPAIEKDFKDGTFDPDNLEKYGFE
PTQEGKLFQLYPRNFLR

Sequence of entity 11 (C), FASTA

>9PR4_11 Translational activator of cytochrome c oxidase 1 (chains C)
MSAWAAASLSRAAARCLLARGPGVRAAPPRDPRPSHPEPRGCGAAPGRTLHFTAAVPAGH
NKWSKVRHIKGPKDVERSRIFSKLCLNIRLAVKEGGPNPEHNSNLANILEVCRSKHMPKS
TIETALKMEKSKDTYLLYEGRGPGGSSLLIEALSNSSHKCQADIRHILNKNGGVMAVGAR
HSFDKKGVIVVEVEDREKKAVNLERALEMAIEAGAEDVKETEDEEERNVFKFICDASSLH
QVRKKLDSLGLCSVSCALEFIPNSKVQLAEPDLEQAAHLIQALSNHEDVIHVYDNIE

Sequence of entity 12 (D), FASTA

>9PR4_12 39S ribosomal protein L2, mitochondrial (chains D)
MALCALTRALRSLNLAPPTVAAPAPSLFPAAQMMNNGLLQQPSALMLLPCRPVLTSVALN
ANFVSWKSRTKYTITPVKMRKSGGRDHTGRIRVHGIGGGHKQRYRMIDFLRFRPEETKSG
PFEEKVIQVRYDPCRSADIALVAGGSRKRWIIATENMQAGDTILNSNHIGRMAVAAREGD
AHPLGALPVGTLINNVESEPGRGAQYIRAAGTCGVLLRKVNGTAIIQLPSKRQMQVLETC
VATVGRVSNVDHNKRVIGKAGRNRWLGKRPNSGRWHRKGGWAGRKIRPLPPMKSYVKLPS
ASAQS

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	2
MG	Magnesium ion	Mg	141
FES	FE2/S2 (inorganic) cluster	Fe2 S2	1
SPD	Spermidine	C7 H19 N3	2
PUT	1,4-diaminobutane	C4 H12 N2	1
VAL	Valine	C5 H11 N O2	1

Water and common crystallization additives (K) are not listed.

Primary citation

Structural basis of TACO1-mediated efficient mitochondrial translation. Wang, S., Brischigliaro, M., Zhang, Y. et al. Nat Commun (2026) 17. DOI 10.1038/s41467-026-69156-y · PubMed

Other PDB entries of the same protein (UniProt Q9BYC8 (AlphaFold model), which also has an AlphaFold model), best resolution first:

7OF0 2.2 Å, Structure of a human mitochondrial ribosome large subunit assembly intermediate in…
7QI4 2.21 Å, Human mitochondrial ribosome at 2.2 A resolution (bound to partly built tRNAs and mRNA)
8RRI 2.4 Å, Human mitochondrial ribosome in complex with antibiotic tigecycline
8QU5 2.42 Å, mt-LSU assembly intermediate in GTPBP8 knock-out cells, state 2
9OLF 2.46 Å, Membrane-associated human mitoribosome in complex with TACO1
7OF7 2.5 Å, Structure of a human mitochondrial ribosome large subunit assembly intermediate in…
7PO4 2.56 Å, Assembly intermediate of human mitochondrial ribosome large subunit (largely unfolded…
6ZM6 2.59 Å, Human mitochondrial ribosome in complex with mRNA, A/A tRNA and P/P tRNA
7O9M 2.6 Å, Human mitochondrial ribosome large subunit assembly intermediate with MTERF4-NSUN4,…
7OF6 2.6 Å, Structure of mature human mitochondrial ribosome large subunit in complex with GTPBP6…
9CN3 2.62 Å, Human 39S mitoribosome in complex with antibiotic Linezolid
7QI5 2.63 Å, Human mitochondrial ribosome in complex with mRNA, A/A-, P/P- and E/E-tRNAs at 2.63 A…

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