9MF0: KRAS(GDP)

Crystal structure of KRAS(GDP) bound to LZTR1(Kelch domain). Determined by X-ray diffraction at 3.3 Å resolution. Released 10 Sept 2025.

Method: X-ray diffraction
Resolution: 3.3 Å
Organism: Homo sapiens
Chains: 10
Atoms: 19,642
Mol. weight: 281.5 kDa
Ligands: GDP, MG
Released: 10 Sept 2025

Explore 9MF0 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9MF0 contains 72 α-helices and 234 β-strands across 10 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 A and G: 7 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	1-10	10	1
α-helix	16-25	10
β-strand	38-45	8	1
β-strand	50-57	8	1
α-helix	64-65	2
α-helix	66-74	9
β-strand	77-83	7	1
α-helix	87-91	5
α-helix	93-104	12
β-strand	111-116	6	1
α-helix	127-137	11
β-strand	141-143	3	1
β-strand	145	1	2
β-strand	150	1	2
α-helix	152-168	17

Chain B: 8 helices, 38 β-strands

Element	Residues	Length	Sheet
β-strand	53-57	5	3
α-helix	58-59	2
β-strand	68	1	3
β-strand	69	1	4
β-strand	72-76	5	4
β-strand	79-86	8	4
β-strand	91-99	9	4
β-strand	104	1	4
β-strand	105-106	2	5
β-strand	107	1	4
α-helix	113-116	4
β-strand	118-119	2	6
β-strand	122-126	5	6
β-strand	129-139	11	6
β-strand	144-156	13	6
β-strand	161-164	4	6
α-helix	172-174	3
β-strand	175	1	6
β-strand	176	1	7
β-strand	179-183	5	7
β-strand	186-193	8	7
β-strand	198-206	9	7
β-strand	215-217	3	7
β-strand	221-222	2	8
α-helix	224-226	3
β-strand	228	1	7
β-strand	229	1	9
β-strand	233-236	4	8
β-strand	239-243	5	8
β-strand	246	1	9
α-helix	248-250	3
β-strand	255-259	5	8
β-strand	264-267	4	8
α-helix	270-275	6
α-helix	277-283	7
β-strand	285-291	7	10
β-strand	295-302	8	10
β-strand	303	1	11
β-strand	306	1	11
β-strand	312-315	4	10
β-strand	320-323	4	10
α-helix	384-387	4
β-strand	389	1	3
β-strand	392-396	5	3
β-strand	399-406	8	3
β-strand	412-420	9	3

Chain C: 7 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	2-10	9	5
α-helix	16-25	10
β-strand	38-45	8	5
β-strand	50-57	8	5
α-helix	64-65	2
α-helix	66-74	9
β-strand	77-83	7	5
α-helix	87-91	5
α-helix	93-104	12
β-strand	111-116	6	5
α-helix	127-137	11
β-strand	141-143	3	5
β-strand	145	1	12
β-strand	150	1	12
α-helix	152-168	17

Chain D: 8 helices, 44 β-strands

Element	Residues	Length	Sheet
β-strand	53-57	5	13
α-helix	58-59	2
β-strand	68	1	14
β-strand	69	1	15
β-strand	72-76	5	15
β-strand	79-86	8	15
β-strand	91-99	9	15
β-strand	104	1	15
β-strand	105-106	2	16
β-strand	107	1	15
α-helix	113-116	4
β-strand	118-119	2	17
β-strand	122-126	5	18
β-strand	129-133	5	18
β-strand	136-139	4	17
β-strand	144-148	5	17
β-strand	149	1	19
β-strand	152-156	5	18
β-strand	161-164	4	18
α-helix	172-174	3
β-strand	175	1	19
β-strand	176	1	20
β-strand	179-183	5	20
β-strand	186-193	8	20
β-strand	198-206	9	20
β-strand	215-217	3	20
β-strand	221-222	2	21
α-helix	224-226	3
β-strand	228	1	20
β-strand	229	1	22
β-strand	233-236	4	21
β-strand	239-243	5	21
β-strand	246	1	22
α-helix	248-250	3
β-strand	255-259	5	21
β-strand	264-267	4	21
α-helix	270-272	3
α-helix	278-283	6
β-strand	285-291	7	23
β-strand	295-302	8	23
β-strand	303	1	24
β-strand	306	1	24
β-strand	312-315	4	23
β-strand	320-323	4	23
α-helix	384-387	4
β-strand	389	1	25
β-strand	392-396	5	13
β-strand	399-403	5	13
β-strand	406	1	25
β-strand	412	1	25
β-strand	413	1	14
β-strand	416-420	5	13

Chain E: 7 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	2-9	8	26
α-helix	16-25	10
β-strand	38-46	9	26
β-strand	49-57	9	26
α-helix	64-65	2
α-helix	66-74	9
β-strand	77-83	7	26
α-helix	87-91	5
α-helix	93-104	12
β-strand	111-116	6	26
α-helix	127-137	11
β-strand	141-143	3	26
β-strand	145	1	27
β-strand	150	1	27
α-helix	152-167	16

Chain F: 8 helices, 39 β-strands

Element	Residues	Length	Sheet
β-strand	53-57	5	28
α-helix	58-59	2
β-strand	68	1	29
β-strand	69	1	30
β-strand	72-76	5	30
β-strand	79-86	8	30
β-strand	91-99	9	30
β-strand	104	1	30
β-strand	105-106	2	1
β-strand	107	1	30
α-helix	113-116	4
β-strand	118-119	2	31
β-strand	122-126	5	31
β-strand	129-139	11	31
β-strand	144-156	13	31
β-strand	161-164	4	31
α-helix	172-174	3
β-strand	175	1	31
β-strand	176	1	32
β-strand	179-183	5	32
β-strand	186-193	8	32
β-strand	198-206	9	32
β-strand	216-217	2	32
β-strand	221-222	2	33
α-helix	224-226	3
β-strand	228	1	32
β-strand	229	1	34
β-strand	233-236	4	33
β-strand	239-243	5	33
β-strand	246	1	34
α-helix	248-250	3
β-strand	255-259	5	33
β-strand	264-267	4	33
α-helix	268-269	2
α-helix	279-283	5
β-strand	285-291	7	35
β-strand	295-302	8	35
β-strand	312-315	4	35
β-strand	320-323	4	35
α-helix	384-387	4
β-strand	389	1	36
β-strand	392-396	5	28
β-strand	399-403	5	28
β-strand	406	1	36
β-strand	412	1	36
β-strand	413	1	29
β-strand	416-420	5	28

Chain H: 7 helices, 39 β-strands

Element	Residues	Length	Sheet
β-strand	53-57	5	38
α-helix	58-59	2
β-strand	68	1	39
β-strand	69	1	40
β-strand	72-76	5	40
β-strand	79-86	8	40
β-strand	91-99	9	40
β-strand	104	1	40
β-strand	105-106	2	41
β-strand	107	1	40
α-helix	113-116	4
β-strand	118-119	2	42
β-strand	122-126	5	42
β-strand	129-139	11	42
β-strand	144-156	13	42
β-strand	161-164	4	42
α-helix	172-174	3
β-strand	175	1	42
β-strand	176	1	43
β-strand	179-183	5	43
β-strand	186-193	8	43
β-strand	198-206	9	43
β-strand	215-217	3	43
β-strand	221-222	2	44
α-helix	224-226	3
β-strand	228	1	43
β-strand	233-236	4	44
β-strand	239-243	5	44
α-helix	248-250	3
β-strand	255-259	5	44
β-strand	264-267	4	44
α-helix	278-283	6
β-strand	285	1	45
β-strand	288-291	4	46
β-strand	295-299	5	46
β-strand	302	1	45
β-strand	312-315	4	46
β-strand	320-323	4	46
α-helix	384-387	4
β-strand	389	1	47
β-strand	392-396	5	38
β-strand	399-403	5	38
β-strand	406	1	47
β-strand	412	1	47
β-strand	413	1	39
β-strand	416-420	5	38

Chain I: 6 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	1-10	10	41
α-helix	16-25	10
β-strand	38-45	8	41
β-strand	50-57	8	41
α-helix	64-65	2
α-helix	66-74	9
β-strand	77-83	7	41
α-helix	87-104	18
β-strand	111-116	6	41
α-helix	127-137	11
β-strand	141-143	3	41
β-strand	145	1	48
β-strand	150	1	48
α-helix	152-168	17

1 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
Isoform 2B of GTPase KRas	A, C, E, G, I	protein	170	Homo sapiens	P01116 (AlphaFold model)
Leucine-zipper-like transcriptional regulator 1	B, D, F, H, J	protein	322	Homo sapiens	Q8N653 (AlphaFold model)

Sequence of entity 1 (A, C, E, G, I), FASTA

>9MF0_1 Isoform 2B of GTPase KRas (chains A, C, E, G, I)
GMTEYKLVVVGAGGVGKSALTIQLIQNHFVDEYDPAIEDSYRKQVVIDGETCLLDILDTA
GQAEYSAMRDQYMRTGEGFLCVFAINNTKSFEDIHHYREQIKRVKDSEDVPMVLVGNKCD
LPSRTVDTKQAQDLARSYGIPFIETSAKTRQGVDDAFYTLVREIRKHKEK

Sequence of entity 2 (B, D, F, H, J), FASTA

>9MF0_2 Leucine-zipper-like transcriptional regulator 1 (chains B, D, F, H, J)
GTVHRWRRLPPCDEFVGARRSKHTVVAYKDAIYVFGGDNGKTMLNDLLRFDVKDCSWCRA
FTTGTPPAPRYHHSAVVYGSSMFVFGGYTGDIYSNSNLKNKNDLFEYKFATGQWTEWKIE
GRLPVARSAHGATVYSDKLWIFAGYDGNARLNDMWTIGLQDRELTCWEEVAQSGEIPPSC
CNFPVAVCRDKMFVFSGQSGAKITNNLFQFEFKDKTWTRIPTEHLLRGSPPPPQRRYGHT
MVAFDRHLYVFGGAADNTLPNELHCYDVDFQTWEVVQPSSDSELPSGRLFHAAAVISDAM
YIFGGTVDNNIRSGEMYRFQFS

Ligands and cofactors

ID	Name	Formula	Copies
GDP	Guanosine-5'-diphosphate	C10 H15 N5 O11 P2	5
MG	Magnesium ion	Mg	5

Primary citation

Structural basis for LZTR1 recognition of RAS GTPases for degradation. Dharmaiah, S., Bonsor, D.A., Mo, S.P. et al. Science (2025) 389:1112-1117. DOI 10.1126/science.adv7088 · PubMed

Other PDB entries of the same protein (UniProt P01116 (AlphaFold model), which also has an AlphaFold model), best resolution first:

9IAY 0.95 Å, Structure of 10 in complex with GDP-KRAS
9IAW 1.0 Å, Structure of 5 in complex with GDP-KRAS
12XE 1.01 Å, Crystal structure of KRAS(Q61K) bound to GppNHp and ligand
8ONV 1.01 Å, KRAS-G13D in complex with BI-2493
6P0Z 1.01 Å, Crystal structure of N-acetylated KRAS (2-169) bound to GDP and Mg
9IB5 1.01 Å, Structure of 18 (BI-2493) in complex with GDP-KRAS
8AZZ 1.02 Å, KRAS-G12V in complex with BI-2865
8AZX 1.04 Å, KRAS-G12C in complex with BI-2865
8B00 1.04 Å, KRAS-G13D in complex with BI-2865
8TVK 1.04 Å, KRAS 1-169 G12C Mutant at 100k
4QL3 1.04 Å, Crystal Structure of a GDP-bound G12R Oncogenic Mutant of Human GTPase KRas
8AZV 1.05 Å, KRAS in complex with BI-2865

Browse structure collections

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