9VPC: IF1 bound bovine ATP synthase tetramer

Cryo-EM structure of the IF1 bound bovine ATP synthase tetramer. Determined by electron microscopy at 7.2 Å resolution. Released 1 Jul 2026.

Method: Electron microscopy
Resolution: 7.2 Å
Organism: Bos taurus
Chains: 116
Atoms: 101,327
Mol. weight: 2788.45 kDa
Released: 1 Jul 2026

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

Secondary structure: helices and β-sheets

9VPC contains 942 α-helices and 692 β-strands across 116 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.

Chains 18 and 38: 4 helices, 0 β-strands

Element	Residues	Length
α-helix	2-4	3
α-helix	9-20	12
α-helix	23-30	8
α-helix	34-38	5

Chains 1a and 3a: 11 helices, 0 β-strands

Element	Residues	Length
α-helix	3-8	6
α-helix	19-30	12
α-helix	41-60	20
α-helix	65-68	4
α-helix	69-85	17
α-helix	94-96	3
α-helix	98-119	22
α-helix	121-128	8
α-helix	135-137	3
α-helix	138-184	47
α-helix	187-225	39

Chains 1A and 3A: 23 helices, 27 β-strands

Element	Residues	Length	Sheet
α-helix	9-17	9
α-helix	19-21	3
β-strand	30	1	1
β-strand	31-34	4	2
β-strand	39-41	3	2
β-strand	51-54	4	2
β-strand	60-66	7	2
β-strand	71-75	5	2
α-helix	79-81	3
β-strand	87	1	1
β-strand	89-94	6	2
β-strand	96-98	3	3
β-strand	107-108	2	4
β-strand	114	1	4
β-strand	126-128	3	3
β-strand	139	1	5
β-strand	145	1	6
α-helix	152-155	4
β-strand	160	1	6
β-strand	164	1	4
β-strand	167-169	3	4
α-helix	175-190	16
α-helix	195-197	3
β-strand	199-208	10	4
α-helix	210-220	11
β-strand	230-235	6	4
α-helix	241-258	18
β-strand	263-269	7	4
α-helix	271-283	13
α-helix	287-289	3
α-helix	291-293	3
α-helix	298-307	10
β-strand	312	1	5
β-strand	322-323	2	4
β-strand	326-328	3	4
α-helix	330-332	3
α-helix	337-345	9
β-strand	348-349	2	7
β-strand	350-352	3	4
α-helix	354-359	6
β-strand	365	1	4
β-strand	371-372	2	7
α-helix	375-377	3
α-helix	381-405	25
α-helix	412-427	16
α-helix	438-449	12
α-helix	452-454	3
α-helix	461-474	14
α-helix	478-486	9
α-helix	491-507	17

Chains 1b and 3b: 4 helices, 2 β-strands

Element	Residues	Length	Sheet
β-strand	12-13	2	168
β-strand	17-18	2	168
α-helix	19-29	11
α-helix	32-48	17
α-helix	55-188	134
α-helix	193-206	14

Chains 1B and 3B: 20 helices, 23 β-strands

Element	Residues	Length	Sheet
β-strand	30-31	2	9
β-strand	35	1	9
β-strand	38-42	5	9
β-strand	54-55	2	10
β-strand	63-66	4	9
β-strand	71-74	4	9
β-strand	87	1	9
β-strand	88-89	2	10
β-strand	96-98	3	11
β-strand	107-108	2	12
β-strand	114	1	12
β-strand	126-128	3	11
α-helix	132-134	3
β-strand	145	1	13
α-helix	146-147	2
α-helix	151-156	6
β-strand	160	1	13
β-strand	166-169	4	14
α-helix	176-189	14
β-strand	199	1	15
β-strand	201-206	6	12
α-helix	210-221	12
β-strand	229-234	6	12
α-helix	240-260	21
β-strand	263-265	3	15
β-strand	268	1	12
α-helix	272-283	12
α-helix	287-289	3
α-helix	291-293	3
α-helix	298-306	9
β-strand	320-322	3	15
β-strand	326-328	3	14
α-helix	330-332	3
α-helix	337-342	6
β-strand	349-352	4	14
α-helix	354-359	6
α-helix	381-400	20
α-helix	412-426	15
α-helix	429-431	3
α-helix	438-450	13
α-helix	461-474	14
α-helix	477-485	9
α-helix	491-507	17

Chains 1C and 3C: 25 helices, 24 β-strands

Element	Residues	Length	Sheet
α-helix	7-18	12
β-strand	27	1	17
β-strand	28-34	7	2
β-strand	38-43	6	2
β-strand	51-55	5	2
β-strand	60-66	7	2
β-strand	71-75	5	2
β-strand	87-94	8	2
β-strand	97-98	2	18
β-strand	107-109	3	19
β-strand	114	1	19
β-strand	126-127	2	18
α-helix	131-134	4
β-strand	139	1	20
β-strand	145	1	21
α-helix	151-155	5
β-strand	160	1	21
β-strand	164	1	22
β-strand	165-169	5	23
α-helix	175-190	16
β-strand	199-206	8	19
α-helix	210-222	13
α-helix	226-228	3
β-strand	230-234	5	19
α-helix	240-259	20
β-strand	263-266	4	19
α-helix	271-284	14
α-helix	287-289	3
α-helix	291-293	3
α-helix	294-295	2
α-helix	298-306	9
β-strand	312	1	20
β-strand	322	1	19
β-strand	323	1	22
β-strand	326-328	3	23
α-helix	330-332	3
α-helix	337-345	9
β-strand	348-351	4	23
α-helix	352-353	2
α-helix	355-358	4
α-helix	375-377	3
α-helix	381-399	19
α-helix	412-426	15
α-helix	435-437	3
α-helix	438-449	12
α-helix	458-460	3
α-helix	461-474	14
α-helix	477-486	10
α-helix	491-507	17

Chains 1d and 3d: 11 helices, 0 β-strands

Element	Residues	Length
α-helix	12-15	4
α-helix	21-23	3
α-helix	24-43	20
α-helix	53-59	7
α-helix	65-75	11
α-helix	88-90	3
α-helix	91-97	7
α-helix	98-105	8
α-helix	106-119	14
α-helix	127-129	3
α-helix	132-138	7

Chains 1D and 3D: 19 helices, 26 β-strands

Element	Residues	Length	Sheet
β-strand	10-17	8	2
β-strand	20-25	6	2
α-helix	29-31	3
β-strand	35-38	4	2
β-strand	46-52	7	2
β-strand	58-62	5	2
β-strand	74-77	4	2
β-strand	83-85	3	24
α-helix	88-90	3
β-strand	95	1	25
β-strand	101	1	25
β-strand	113-115	3	24
α-helix	120-122	3
β-strand	132	1	26
α-helix	138-143	6
β-strand	147	1	26
β-strand	153	1	27
β-strand	154	1	28
α-helix	163-176	14
β-strand	181-188	8	25
α-helix	190-203	14
β-strand	215-221	7	25
α-helix	226-245	20
β-strand	250-255	6	25
α-helix	258-270	13
β-strand	277	1	29
β-strand	281	1	29
α-helix	285-293	9
β-strand	303-308	6	25
β-strand	309	1	28
α-helix	313-315	3
α-helix	320-328	9
β-strand	331	1	30
β-strand	332	1	27
β-strand	335	1	31
α-helix	337-342	6
β-strand	348	1	31
β-strand	355	1	30
α-helix	360-363	4
α-helix	365-387	23
α-helix	398-413	16
α-helix	422-425	4
α-helix	434-446	13
α-helix	455-457	3
α-helix	463-476	14

49 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
ATP synthase F(0) complex subunit 8	18, 28, 38, 48	protein	66	Bos taurus	P03929 (AlphaFold model)
ATP synthase subunit alpha	1A, 1B, 1C, 2A, 2B, 2C, 3A, 3B, 3C, 4A, 4B, 4C	protein	553	Bos taurus	F1MLB8 (AlphaFold model)
ATP synthase F(1) complex catalytic subunit beta, mitochondrial	1D, 1E, 1F, 2D, 2E, 2F, 3D, 3E, 3F, 4D, 4E, 4F	protein	528	Bos taurus	P00829 (AlphaFold model)
ATP synthase F(1) complex subunit gamma, mitochondrial	1G, 2G, 3G, 4G	protein	298	Bos taurus	P05631 (AlphaFold model)
ATP synthase F(1) complex subunit delta, mitochondrial	1H, 2H, 3H, 4H	protein	168	Bos taurus	P05630
ATP synthase F(1) complex subunit epsilon, mitochondrial	1I, 2I, 3I, 4I	protein	51	Bos taurus	P05632
ATPase inhibitor, mitochondrial	1J, 2J, 3J, 4J	protein	109	Bos taurus	P01096
ATP synthase F(0) complex subunit C2, mitochondrial	1K, 1L, 1M, 1N, 1O, 1P, 1Q, 1R, 2K, 2L, 2M, 2N, 2O, 2P, 2Q, 2R, 3K, 3L, 3M, 3N, 3O, 3P, 3Q, 3R, 4K, 4L, 4M, 4N, 4O, 4P, 4Q, 4R	protein	143	Bos taurus	P07926
ATP synthase peripheral stalk subunit OSCP, mitochondrial	1S, 2S, 3S, 4S	protein	213	Bos taurus	P13621
ATP synthase F(0) complex subunit a	1a, 2a, 3a, 4a	protein	226	Bos taurus	P00847
ATP synthase peripheral stalk subunit b, mitochondrial	1b, 2b, 3b, 4b	protein	256	Bos taurus	P13619
ATP synthase peripheral stalk subunit d, mitochondrial	1d, 2d, 3d, 4d	protein	161	Bos taurus	P13620

6 more molecules are not listed.

Sequence of entity 1 (18, 28, 38, 48), FASTA

>9VPC_1 ATP synthase F(0) complex subunit 8 (chains 18, 28, 38, 48)
MPQLDTSTWLTMILSMFLTLFIIFQLKVSKHNFYHNPELTPTKMLKQNTPWETKWTKIYL
PLLLPL

Sequence of entity 2 (1A, 1B, 1C, 2A, 2B, 2C, 3A, 3B, 3C, 4A, 4B, 4C), FASTA

>9VPC_2 ATP synthase subunit alpha (chains 1A, 1B, 1C, 2A, 2B, 2C, 3A, 3B, 3C, 4A, 4B, 4C)
MLSVRVAAAVARALPRRAGLVSKNALGSSFIAARNLHASNSRLQKTGTAEVSSILEERIL
GADTSVDLEETGRVLSIGDGIARVHGLRNVQAEEMVEFSSGLKGMSLNLEPDNVGVVVFG
NDKLIKEGDIVKRTGAIVDVPVGEELLGRVVDALGNAIDGKGPIGSKARRRVGLKAPGII
PRISVREPMQTGIKAVDSLVPIGRGQRELIIGDRQTGKTSIAIDTIINQKRFNDGTDEKK
KLYCIYVAIGQKRSTVAQLVKRLTDADAMKYTIVVSATASDAAPLQYLAPYSGCSMGEYF
RDNGKHALIIYDDLSKQAVAYRQMSLLLRRPPGREAYPGDVFYLHSRLLERAAKMNDAFG
GGSLTALPVIETQAGDVSAYIPTNVISITDGQIFLETELFYKGIRPAINVGLSVSRVGSA
AQTRAMKQVAGTMKLELAQYREVAAFAQFGSDLDAATQQLLSRGVRLTELLKQGQYSPMA
IEEQVAVIYAGVRGYLDKLEPSKITKFENAFLSHVISQHQALLGKIRTDGKISEESDAKL
KEIVTNFLAGFEA

Sequence of entity 3 (1D, 1E, 1F, 2D, 2E, 2F, 3D, 3E, 3F, 4D, 4E, 4F), FASTA

>9VPC_3 ATP synthase F(1) complex catalytic subunit beta, mitochondrial (chains 1D, 1E, 1F, 2D, 2E, 2F, 3D, 3E, 3F, 4D, 4E, 4F)
MLGLVGRVVAASASGALRGLSPSAPLPQAQLLLRAAPAALQPARDYAAQASPSPKAGATT
GRIVAVIGAVVDVQFDEGLPPILNALEVQGRETRLVLEVAQHLGESTVRTIAMDGTEGLV
RGQKVLDSGAPIRIPVGPETLGRIMNVIGEPIDERGPIKTKQFAAIHAEAPEFVEMSVEQ
EILVTGIKVVDLLAPYAKGGKIGLFGGAGVGKTVLIMELINNVAKAHGGYSVFAGVGERT
REGNDLYHEMIESGVINLKDATSKVALVYGQMNEPPGARARVALTGLTVAEYFRDQEGQD
VLLFIDNIFRFTQAGSEVSALLGRIPSAVGYQPTLATDMGTMQERITTTKKGSITSVQAI
YVPADDLTDPAPATTFAHLDATTVLSRAIAELGIYPAVDPLDSTSRIMDPNIVGSEHYDV
ARGVQKILQDYKSLQDIIAILGMDELSEEDKLTVSRARKIQRFLSQPFQVAEVFTGHLGK
LVPLKETIKGFQQILAGEYDHLPEQAFYMVGPIEEAVAKADKLAEEHS

Sequence of entity 4 (1G, 2G, 3G, 4G), FASTA

>9VPC_4 ATP synthase F(1) complex subunit gamma, mitochondrial (chains 1G, 2G, 3G, 4G)
MFSRAGVAGLSAWTVQPQWIQVRNMATLKDITRRLKSIKNIQKITKSMKMVAAAKYARAE
RELKPARVYGVGSLALYEKADIKTPEDKKKHLIIGVSSDRGLCGAIHSSVAKQMKSEAAN
LAAAGKEVKIIGVGDKIRSILHRTHSDQFLVTFKEVGRRPPTFGDASVIALELLNSGYEF
DEGSIIFNRFRSVISYKTEEKPIFSLDTISSAESMSIYDDIDADVLRNYQEYSLANIIYY
SLKESTTSEQSARMTAMDNASKNASEMIDKLTLTFNRTRQAVITKELIEIISGAAALD

Sequence of entity 5 (1H, 2H, 3H, 4H), FASTA

>9VPC_5 ATP synthase F(1) complex subunit delta, mitochondrial (chains 1H, 2H, 3H, 4H)
MLPSALLRRPGLGRLVRQVRLYAEAAAAQAPAAGPGQMSFTFASPTQVFFNSANVRQVDV
PTQTGAFGILAAHVPTLQVLRPGLVVVHAEDGTTSKYFVSSGSVTVNADSSVQLLAEEAV
TLDMLDLGAAKANLEKAQSELLGAADEATRAEIQIRIEANEALVKALE

Sequence of entity 6 (1I, 2I, 3I, 4I), FASTA

>9VPC_6 ATP synthase F(1) complex subunit epsilon, mitochondrial (chains 1I, 2I, 3I, 4I)
MVAYWRQAGLSYIRYSQICAKAVRDALKTEFKANAMKTSGSTIKIVKVKKE

Sequence of entity 7 (1J, 2J, 3J, 4J), FASTA

>9VPC_7 ATPase inhibitor, mitochondrial (chains 1J, 2J, 3J, 4J)
MAATALAARTRQAVWSVWAMQGRGFGSESGDNVRSSAGAVRDAGGAFGKREQAEEERYFR
ARAKEQLAALKKHHENEISHHAKEIERLQKEIERHKQSIKKLKQSEDDD

Sequence of entity 8 (1K, 1L, 1M, 1N, 1O, 1P, 1Q, 1R, 2K, 2L, 2M, 2N, 2O, 2P, 2Q, 2R, 3K, 3L, 3M, 3N, 3O, 3P, 3Q, 3R, 4K, 4L, 4M, 4N, 4O, 4P, 4Q, 4R), FASTA

>9VPC_8 ATP synthase F(0) complex subunit C2, mitochondrial (chains 1K, 1L, 1M, 1N, 1O, 1P, 1Q, 1R, 2K, 2L, 2M, 2N, 2O, 2P, 2Q, 2R, 3K, 3L, 3M, 3N, 3O, 3P, 3Q, 3R, 4K, 4L, 4M, 4N, 4O, 4P, 4Q, 4R)
MYTCAKFVSTPSLIRRTSTVLSRSLSAVVVRRPETLTDESHSSLAVVPRPLTTSLTPSRS
FQTSAISRDIDTAAKFIGAGAATVGVAGSGAGIGTVFGSLIIGYARNPSLKQQLFSYAIL
GFALSEAMGLFCLMVAFLILFAM

Sequence of entity 9 (1S, 2S, 3S, 4S), FASTA

>9VPC_9 ATP synthase peripheral stalk subunit OSCP, mitochondrial (chains 1S, 2S, 3S, 4S)
MAALAVSGLSQQVRCFSTSVVRPFAKLVRPPVQIYGIEGRYATALYSAASKQNKLEQVEK
ELLRVGQILKEPKMAASLLNPYVKRSVKVKSLSDMTAKEKFSPLTSNLINLLAENGRLTN
TPAVISAFSTMMSVHRGEVPCTVTTASALDEATLTELKTVLKSFLSKGQVLKLEVKIDPS
IMGGMIVRIGEKYVDMSAKTKIQKLSRAMREIL

Sequence of entity 10 (1a, 2a, 3a, 4a), FASTA

>9VPC_10 ATP synthase F(0) complex subunit a (chains 1a, 2a, 3a, 4a)
MNENLFTSFITPVILGLPLVTLIVLFPSLLFPTSNRLVSNRFVTLQQWMLQLVSKQMMSI
HNSKGQTWTLMLMSLILFIGSTNLLGLLPHSFTPTTQLSMNLGMAIPLWAGAVITGFRNK
TKASLAHFLPQGTPTPLIPMLVIIETISLFIQPMALAVRLTANITAGHLLIHLIGGATLA
LMSISTTTALITFTILILLTILEFAVAMIQAYVFTLLVSLYLHDNT

Sequence of entity 11 (1b, 2b, 3b, 4b), FASTA

>9VPC_11 ATP synthase peripheral stalk subunit b, mitochondrial (chains 1b, 2b, 3b, 4b)
MLSRVVLSAAAAAAPSLKNAALLGPGVLQATRIFHTGQPSLAPVPPLPEHGGKVRFGLIP
EEFFQFLYPKTGVTGPYVLGTGLILYLLSKEIYVITPETFSAISTIGFLVYIVKKYGASV
GEFADKLNEQKIAQLEEVKQASIKQIQDAIDMEKSQQALVQKRHYLFDVQRNNIAMALEV
TYRERLHRVYREVKNRLDYHISVQNMMRQKEQEHMINWVEKRVVQSISAQQEKETIAKCI
ADLKLLSKKAQAQPVM

Sequence of entity 12 (1d, 2d, 3d, 4d), FASTA

>9VPC_12 ATP synthase peripheral stalk subunit d, mitochondrial (chains 1d, 2d, 3d, 4d)
MAGRKLALKTIDWVAFGEIIPRNQKAVANSLKSWNETLTSRLATLPEKPPAIDWAYYKAN
VAKAGLVDDFEKKFNALKVPIPEDKYTAQVDAEEKEDVKSCAEFLTQSKTRIQEYEKELE
KMRNIIPFDQMTIEDLNEVFPETKLDKKKYPYWPHRPIETL

Primary citation

A planar dimer of bovine ATP synthase. Jiko, C., Nakano, A., Teshirogi, Y. et al. Cell Death Differ (2026). DOI 10.1038/s41418-026-01797-4 · PubMed

Other PDB entries of the same protein (UniProt P03929 (AlphaFold model), which also has an AlphaFold model), best resolution first:

6ZQM 3.29 Å, bovine ATP synthase monomer state 2 (combined)
9W2R 3.4 Å, Cryo-EM structure of FoF1-ATPase monomer state 1 on the bovine heart submitochondrial…
6ZIT 3.49 Å, bovine ATP synthase Stator domain, state 2
6ZBB 3.61 Å, bovine ATP synthase Fo domain
6ZPO 4.0 Å, bovine ATP synthase monomer state 1 (combined)
6ZQN 4.0 Å, bovine ATP synthase monomer state 3 (combined)
9W2S 4.0 Å, Cryo-EM structure of FoF1-ATPase monomer state 3 on the bovine heart submitochondrial…
9W2T 4.1 Å, Cryo-EM structure of Fo domain of FoF1-ATPase monomer state on the bovine heart…
6ZIQ 4.33 Å, bovine ATP synthase stator domain, state 1
9VPB 5.0 Å, Cryo-EM structure of the IF1 bound bovine F-ATP synthase planar dimer
6ZIU 6.02 Å, bovine ATP synthase stator domain, state 3
7AJF 8.45 Å, bovine ATP synthase dimer state2:state2

Browse structure collections

About this viewer

MolViewer shows 9VPC directly in your browser with nothing to install. Switch between cartoon, ball-and-stick, spacefill and surface views, color by chain, secondary structure or B-factor, measure distances, angles and dihedrals, and share or embed the view.