PDB-1pcq: Crystal structure of groEL-groES

ID or keywords:

Entry

Database: PDB / ID: 1pcq

Title

Crystal structure of groEL-groES

Components

groEL proteinGroEL
groES proteinGroEL

Keywords

CHAPERONE

Function / homology

Function and homology information

GroEL-GroES complex /

chaperonin ATPase /

virion assembly / chaperone cofactor-dependent protein refolding / protein folding chaperone /

isomerase activity / ATP-dependent protein folding chaperone / response to radiation / unfolded protein binding /

protein folding ...
Similarity search - Function

Biological species

Escherichia coli (E. coli)

Method

X-RAY DIFFRACTION /

SYNCHROTRON / Resolution: 2.808 Å

Authors

Chaudhry, C. / Farr, G.W. / Todd, M.J. / Rye, H.S. / Brunger, A.T. / Adams, P.D. / Horwich, A.L. / Sigler, P.B.

Citation

Journal: Embo J. / Year: 2003
Title: Role of the gamma-phosphate of ATP in triggering protein folding by GroEL-GroES: function, structure and energetics.
Authors: Chaudhry, C. / Farr, G.W. / Todd, M.J. / Rye, H.S. / Brunger, A.T. / Adams, P.D. / Horwich, A.L. / Sigler, P.B.

History

Deposition	May 16, 2003	Deposition site: RCSB / Processing site: RCSB
Revision 1.0	Oct 14, 2003	Provider: repository / Type: Initial release
Revision 1.1	Apr 29, 2008	Group: Version format compliance
Revision 1.2	Jul 13, 2011	Group: Version format compliance
Revision 1.3	Feb 14, 2024	Group: Data collection / Database references ...Data collection / Database references / Derived calculations / Refinement description Category: chem_comp_atom / chem_comp_bond ...chem_comp_atom / chem_comp_bond / database_2 / pdbx_struct_conn_angle / struct_conn / struct_conn_type / struct_ncs_dom_lim / struct_site Item: _database_2.pdbx_DOI / _database_2.pdbx_database_accession ..._database_2.pdbx_DOI / _database_2.pdbx_database_accession / _struct_conn.conn_type_id / _struct_conn.id / _struct_conn.pdbx_dist_value / _struct_conn.pdbx_leaving_atom_flag / _struct_conn.ptnr1_auth_comp_id / _struct_conn.ptnr1_auth_seq_id / _struct_conn.ptnr1_label_asym_id / _struct_conn.ptnr1_label_atom_id / _struct_conn.ptnr1_label_comp_id / _struct_conn.ptnr1_label_seq_id / _struct_conn.ptnr2_auth_comp_id / _struct_conn.ptnr2_auth_seq_id / _struct_conn.ptnr2_label_asym_id / _struct_conn.ptnr2_label_atom_id / _struct_conn.ptnr2_label_comp_id / _struct_conn.ptnr2_label_seq_id / _struct_conn_type.id / _struct_ncs_dom_lim.beg_auth_comp_id / _struct_ncs_dom_lim.end_auth_comp_id / _struct_ncs_dom_lim.end_label_asym_id / _struct_ncs_dom_lim.end_label_seq_id / _struct_site.pdbx_auth_asym_id / _struct_site.pdbx_auth_comp_id / _struct_site.pdbx_auth_seq_id

Structure viewer	Molecule: MolmilJmol/JSmol

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Download

PDBx/mmCIF format	1pcq.cif.gz	1.3 MB	Display	PDBx/mmCIF format
PDB format	pdb1pcq.ent.gz	1.1 MB	Display	PDB format
PDBx/mmJSON format	1pcq.json.gz		Tree view	PDBx/mmJSON format
Others	Other downloads

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Validation report

Arichive directory	https://data.pdbj.org/pub/pdb/validation_reports/pc/1pcq ftp://data.pdbj.org/pub/pdb/validation_reports/pc/1pcq	HTTPS FTP

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Related structure data

Related structure data	1pf9C C: citing same article (ref.)
Similar structure data	3zq0-d 3zq1-d 1pf9-d 2c7c-d 1sx4-d 1svt-d 1gru-d 4v4o-2 4v4o-1 3zpz-d Search similar-shape structures by Omokage search (details) Similarity search - Function & homologyF&H Search Similarity search - FunctionF&H Search Similarity search - HomologyF&H Search

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Links

PDB pages	PDBj / wwPDB / NCBI
Related items in Molecule of the Month	#32 - Aug 2002 Chaperones similarity (19) #148 - Apr 2012 Ras Protein similarity (1) #80 - Aug 2006 AAA+ Proteases similarity (1) #13 - Jan 2001 Alcohol Dehydrogenase similarity (1)

Deposited unit

A: groEL protein

B: groEL protein

C: groEL protein

D: groEL protein

E: groEL protein

F: groEL protein

G: groEL protein

H: groEL protein

I: groEL protein

J: groEL protein

K: groEL protein

L: groEL protein

M: groEL protein

N: groEL protein

O: groES protein

P: groES protein

Q: groES protein

R: groES protein

S: groES protein

T: groES protein

U: groES protein

hetero molecules

850 kDa, 21 polymers
Search similar-shape structures of this assembly by Omokage search (details)

1

Idetical with deposited unit
defined by author&software

Unit cell

Length a, b, c (Å)	255.546, 266.855, 187.049
Angle α, β, γ (deg.)	90.000, 90.000, 90.000
Int Tables number	18
Space group name H-M	P2₁2₁2

Noncrystallographic symmetry (NCS)

NCS domain:
NCS domain segments:

Refine code: 1

Dom-ID	Component-ID	Ens-ID	Beg auth comp-ID	Beg label comp-ID	End auth comp-ID	End label comp-ID	Auth asym-ID	Label asym-ID	Auth seq-ID	Label seq-ID
1	1	1	ALA	ALA	SER	SER	A	A	2 - 135	1 - 134
2	1	1	ALA	ALA	SER	SER	B	B	2 - 135	1 - 134
3	1	1	ALA	ALA	SER	SER	C	C	2 - 135	1 - 134
4	1	1	ALA	ALA	SER	SER	D	D	2 - 135	1 - 134
5	1	1	ALA	ALA	SER	SER	E	E	2 - 135	1 - 134
6	1	1	ALA	ALA	SER	SER	F	F	2 - 135	1 - 134
7	1	1	ALA	ALA	SER	SER	G	G	2 - 135	1 - 134
8	2	1	GLY	GLY	K	K	A	A - W	410 - 603	409
9	2	1	GLY	GLY	K	K	B	B - AA	410 - 703	409
10	2	1	GLY	GLY	K	K	C	C - EA	410 - 803	409
11	2	1	GLY	GLY	K	K	D	D - IA	410 - 903	409
12	2	1	GLY	GLY	K	K	E	E - MA	410 - 1003	409
13	2	1	GLY	GLY	K	K	F	F - QA	410 - 1103	409
14	2	1	GLY	GLY	K	K	G	G - UA	410 - 1203	409
1	1	2	VAL	VAL	VAL	VAL	A	A	136 - 190	135 - 189
2	1	2	VAL	VAL	VAL	VAL	B	B	136 - 190	135 - 189
3	1	2	VAL	VAL	VAL	VAL	C	C	136 - 190	135 - 189
4	1	2	VAL	VAL	VAL	VAL	D	D	136 - 190	135 - 189
5	1	2	VAL	VAL	VAL	VAL	E	E	136 - 190	135 - 189
6	1	2	VAL	VAL	VAL	VAL	F	F	136 - 190	135 - 189
7	1	2	VAL	VAL	VAL	VAL	G	G	136 - 190	135 - 189
8	2	2	GLY	GLY	GLU	GLU	A	A	375 - 409	374 - 408
9	2	2	GLY	GLY	GLU	GLU	B	B	375 - 409	374 - 408
10	2	2	GLY	GLY	GLU	GLU	C	C	375 - 409	374 - 408
11	2	2	GLY	GLY	GLU	GLU	D	D	375 - 409	374 - 408
12	2	2	GLY	GLY	GLU	GLU	E	E	375 - 409	374 - 408
13	2	2	GLY	GLY	GLU	GLU	F	F	375 - 409	374 - 408
14	2	2	GLY	GLY	GLU	GLU	G	G	375 - 409	374 - 408
1	1	3	GLU	GLU	GLY	GLY	A	A	191 - 374	190 - 373
2	1	3	GLU	GLU	GLY	GLY	B	B	191 - 374	190 - 373
3	1	3	GLU	GLU	GLY	GLY	C	C	191 - 374	190 - 373
4	1	3	GLU	GLU	GLY	GLY	D	D	191 - 374	190 - 373
5	1	3	GLU	GLU	GLY	GLY	E	E	191 - 374	190 - 373
6	1	3	GLU	GLU	GLY	GLY	F	F	191 - 374	190 - 373
7	1	3	GLU	GLU	GLY	GLY	G	G	191 - 374	190 - 373
1	1	4	ALA	ALA	SER	SER	H	H	2 - 135	1 - 134
2	1	4	ALA	ALA	SER	SER	I	I	2 - 135	1 - 134
3	1	4	ALA	ALA	SER	SER	J	J	2 - 135	1 - 134
4	1	4	ALA	ALA	SER	SER	K	K	2 - 135	1 - 134
5	1	4	ALA	ALA	SER	SER	L	L	2 - 135	1 - 134
6	1	4	ALA	ALA	SER	SER	M	M	2 - 135	1 - 134
7	1	4	ALA	ALA	SER	SER	N	N	2 - 135	1 - 134
8	2	4	GLY	GLY	PRO	PRO	H	H	410 - 525	409 - 524
9	2	4	GLY	GLY	PRO	PRO	I	I	410 - 525	409 - 524
10	2	4	GLY	GLY	PRO	PRO	J	J	410 - 525	409 - 524
11	2	4	GLY	GLY	PRO	PRO	K	K	410 - 525	409 - 524
12	2	4	GLY	GLY	PRO	PRO	L	L	410 - 525	409 - 524
13	2	4	GLY	GLY	PRO	PRO	M	M	410 - 525	409 - 524
14	2	4	GLY	GLY	PRO	PRO	N	N	410 - 525	409 - 524
1	1	5	VAL	VAL	VAL	VAL	H	H	136 - 190	135 - 189
2	1	5	VAL	VAL	VAL	VAL	I	I	136 - 190	135 - 189
3	1	5	VAL	VAL	VAL	VAL	J	J	136 - 190	135 - 189
4	1	5	VAL	VAL	VAL	VAL	K	K	136 - 190	135 - 189
5	1	5	VAL	VAL	VAL	VAL	L	L	136 - 190	135 - 189
6	1	5	VAL	VAL	VAL	VAL	M	M	136 - 190	135 - 189
7	1	5	VAL	VAL	VAL	VAL	N	N	136 - 190	135 - 189
8	2	5	GLY	GLY	GLU	GLU	H	H	375 - 409	374 - 408
9	2	5	GLY	GLY	GLU	GLU	I	I	375 - 409	374 - 408
10	2	5	GLY	GLY	GLU	GLU	J	J	375 - 409	374 - 408
11	2	5	GLY	GLY	GLU	GLU	K	K	375 - 409	374 - 408
12	2	5	GLY	GLY	GLU	GLU	L	L	375 - 409	374 - 408
13	2	5	GLY	GLY	GLU	GLU	M	M	375 - 409	374 - 408
14	2	5	GLY	GLY	GLU	GLU	N	N	375 - 409	374 - 408
1	1	6	GLU	GLU	GLY	GLY	H	H	191 - 374	190 - 373
2	1	6	GLU	GLU	GLY	GLY	I	I	191 - 374	190 - 373
3	1	6	GLU	GLU	GLY	GLY	J	J	191 - 374	190 - 373
4	1	6	GLU	GLU	GLY	GLY	K	K	191 - 374	190 - 373
5	1	6	GLU	GLU	GLY	GLY	L	L	191 - 374	190 - 373
6	1	6	GLU	GLU	GLY	GLY	M	M	191 - 374	190 - 373
7	1	6	GLU	GLU	GLY	GLY	N	N	191 - 374	190 - 373
1	1	7	MET	MET	ALA	ALA	O	O	1 - 97	1 - 97
2	1	7	MET	MET	ALA	ALA	P	P	1 - 97	1 - 97
3	1	7	MET	MET	ALA	ALA	Q	Q	1 - 97	1 - 97
4	1	7	MET	MET	ALA	ALA	R	R	1 - 97	1 - 97
5	1	7	MET	MET	ALA	ALA	S	S	1 - 97	1 - 97
6	1	7	MET	MET	ALA	ALA	T	T	1 - 97	1 - 97
7	1	7	MET	MET	ALA	ALA	U	U	1 - 97	1 - 97

NCS ensembles :

ID
1
2
3
4
5
6
7

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Protein , 2 types, 21 molecules ABCDEFGHIJKLMNOPQRSTU

#1: Protein	groEL protein / GroEL / Protein Cpn60 / 60 kDa chaperonin Mass: 55220.105 Da / Num. of mol.: 14 Source method: isolated from a genetically manipulated source Source: (gene. exp.) Escherichia coli (E. coli) Gene: GROL OR GROEL OR MOPA OR B4143 OR C5227 OR Z5748 OR ECS5124 OR SF4297 Plasmid: IQ-TRC / Production host: Escherichia coli (E. coli) / Strain (production host): DH 5 alpha / References: UniProt: P0A6F5
#2: Protein	groES protein / GroEL / Protein Cpn10 / 10 kDa chaperonin Mass: 10400.938 Da / Num. of mol.: 7 Source method: isolated from a genetically manipulated source Source: (gene. exp.) Escherichia coli (E. coli) Gene: GROS OR GROES OR MOPB OR B4142 OR C5226 OR Z5747 OR ECS5123 OR SF4296 Plasmid: IQ-TRC / Production host: Escherichia coli (E. coli) / Strain (production host): DH 5 alpha / References: UniProt: P0A6F9

#1: Protein

groEL protein /

GroEL / Protein Cpn60 / 60 kDa chaperonin

Mass: 55220.105 Da / Num. of mol.: 14
Source method: isolated from a genetically manipulated source
Source: (gene. exp.)

Escherichia coli (E. coli)
Gene: GROL OR GROEL OR MOPA OR B4143 OR C5227 OR Z5748 OR ECS5124 OR SF4297
Plasmid: IQ-TRC / Production host:

Escherichia coli (E. coli) / Strain (production host): DH 5 alpha / References: UniProt: P0A6F5

#2: Protein

groES protein /

GroEL / Protein Cpn10 / 10 kDa chaperonin

Mass: 10400.938 Da / Num. of mol.: 7
Source method: isolated from a genetically manipulated source
Source: (gene. exp.)

Escherichia coli (E. coli)
Gene: GROS OR GROES OR MOPB OR B4142 OR C5226 OR Z5747 OR ECS5123 OR SF4296
Plasmid: IQ-TRC / Production host:

Escherichia coli (E. coli) / Strain (production host): DH 5 alpha / References: UniProt: P0A6F9

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Non-polymers , 4 types, 28 molecules

#3: Chemical	ChemComp-MG / MAGNESIUM ION Mass: 24.305 Da / Num. of mol.: 7 / Source method: obtained synthetically / Formula: Mg
#4: Chemical	ChemComp-K / POTASSIUM ION Mass: 39.098 Da / Num. of mol.: 7 / Source method: obtained synthetically / Formula: K
#5: Chemical	ChemComp-ADP / ADENOSINE-5'-DIPHOSPHATE / Adenosine diphosphate Mass: 427.201 Da / Num. of mol.: 7 / Source method: obtained synthetically / Formula: C₁₀H₁₅N₅O₁₀P₂ / Comment: ADP, energy-carrying molecule*YM
#6: Chemical	ChemComp-AF3 / ALUMINUM FLUORIDE / Aluminium fluoride Mass: 83.977 Da / Num. of mol.: 7 / Source method: obtained synthetically / Formula: AlF₃

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Experiment

Experiment	Method: X-RAY DIFFRACTION / Number of used crystals: 1

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Sample preparation

Crystal

Density Matthews: 3.77 Å³/Da / Density % sol: 67.36 %

Crystal grow

*PLUS

Temperature: 18 ℃ / pH: 5.5 / Method: vapor diffusion, hanging drop / Details: used to seeding, Xu, Z., (1997) Nature, 388, 741.

Components of the solutions

*PLUS

ID	Conc.	Common name	Crystal-ID	Sol-ID
1	12 %	PEG3000	1	reservoir
2	0.25 M	sodium glutamate	1	reservoir
3	100 mM	cacodylate	1	reservoir
4	6 %	PEG3000	1	drop
5	0.125 M	sodium glutamate	1	drop
6	50 mM	cacodylate	1	drop
7	7.5 mg/ml	protein	1	drop

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Data collection

Diffraction	Mean temperature: 100 K
Diffraction source	Source: SYNCHROTRON / Site: APS / Beamline: 19-ID / Wavelength: 1.013
Detector	Type: SBC / Detector: CCD
Radiation	Protocol: SINGLE WAVELENGTH / Monochromatic (M) / Laue (L): M / Scattering type: x-ray
Radiation wavelength	Wavelength: 1.013 Å / Relative weight: 1
Reflection	Resolution: 2.8→50 Å / Num. obs: 278591
Reflection	PLUS Num. obs: 278877 / % possible obs: 90.3 % / Redundancy: 4 % / R_merge(I) obs*: 0.138
Reflection shell	PLUS Highest resolution: 2.8 Å / Lowest resolution: 2.87 Å / % possible obs: 55 % / Redundancy: 3.9 % / R_merge(I) obs: 0.601 / Mean I/σ(I) obs*: 1.3

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Processing

Software

Name	Version	Classification
REFMAC	refmac_5.1.24 24/04/2001	refinement
DENZO		data reduction
SCALEPACK		data scaling

Refinement

Resolution: 2.808→49.386 Å / TLS residual ADP flag: UNVERIFIED / σ(F): 0

	R_factor	Num. reflection	% reflection
R_free	0.2777	5517	1.98 %
R_work	0.2619	-	-
obs	-	278591	-

Solvent computation

Ion probe radii: 0.8 Å / Shrinkage radii: 0.8 Å / VDW probe radii: 1.4 Å

Displacement parameters

B_iso mean: 2.53 Å²

	B_aniso -1	B_aniso -2	B_aniso -3
1-	-2.386 Å²	0 Å²	0 Å²
2-	-	-2.272 Å²	0 Å²
3-	-	-	4.659 Å²

Refinement step

Cycle: LAST / Resolution: 2.808→49.386 Å

	Protein	Nucleic acid	Ligand	Solvent	Total
Num. atoms	59073	0	231	0	59304

Refine LS restraints NCS

Refine-ID: X-RAY DIFFRACTION

Ens-ID	Dom-ID	Auth asym-ID	Number	Type	Rms dev position (Å)	Weight position
1	1	A	1843	tight positional	0.052	0.05
1	1	A	1843	tight thermal	0.087	0.5
1	2	B	1843	tight positional	0.044	0.05
1	2	B	1843	tight thermal	0.07	0.5
1	3	C	1843	tight positional	0.047	0.05
1	3	C	1843	tight thermal	0.077	0.5
1	4	D	1843	tight positional	0.05	0.05
1	4	D	1843	tight thermal	0.082	0.5
1	5	E	1843	tight positional	0.041	0.05
1	5	E	1843	tight thermal	0.072	0.5
1	6	F	1843	tight positional	0.04	0.05
1	6	F	1843	tight thermal	0.071	0.5
1	7	G	1843	tight positional	0.05	0.05
1	7	G	1843	tight thermal	0.075	0.5
2	1	A	647	tight positional	0.037	0.05
2	1	A	647	tight thermal	0.05	0.5
2	2	B	647	tight positional	0.03	0.05
2	2	B	647	tight thermal	0.041	0.5
2	3	C	647	tight positional	0.033	0.05
2	3	C	647	tight thermal	0.046	0.5
2	4	D	647	tight positional	0.034	0.05
2	4	D	647	tight thermal	0.044	0.5
2	5	E	647	tight positional	0.028	0.05
2	5	E	647	tight thermal	0.039	0.5
2	6	F	647	tight positional	0.027	0.05
2	6	F	647	tight thermal	0.044	0.5
2	7	G	647	tight positional	0.031	0.05
2	7	G	647	tight thermal	0.039	0.5
3	1	A	1398	tight positional	0.028	0.05
3	1	A	1398	tight thermal	0.01	0.5
3	2	B	1398	tight positional	0.037	0.05
3	2	B	1398	tight thermal	0.009	0.5
3	3	C	1398	tight positional	0.028	0.05
3	3	C	1398	tight thermal	0.009	0.5
3	4	D	1398	tight positional	0.027	0.05
3	4	D	1398	tight thermal	0.01	0.5
3	5	E	1398	tight positional	0.024	0.05
3	5	E	1398	tight thermal	0.009	0.5
3	6	F	1398	tight positional	0.027	0.05
3	6	F	1398	tight thermal	0.008	0.5
3	7	G	1398	tight positional	0.03	0.05
3	7	G	1398	tight thermal	0.009	0.5
4	1	H	1811	tight positional	0.046	0.05
4	1	H	1811	tight thermal	0.076	0.5
4	2	I	1811	tight positional	0.053	0.05
4	2	I	1811	tight thermal	0.082	0.5
4	3	J	1811	tight positional	0.05	0.05
4	3	J	1811	tight thermal	0.079	0.5
4	4	K	1811	tight positional	0.044	0.05
4	4	K	1811	tight thermal	0.078	0.5
4	5	L	1811	tight positional	0.044	0.05
4	5	L	1811	tight thermal	0.076	0.5
4	6	M	1811	tight positional	0.071	0.05
4	6	M	1811	tight thermal	0.146	0.5
4	7	N	1811	tight positional	0.048	0.05
4	7	N	1811	tight thermal	0.075	0.5
5	1	H	647	tight positional	0.021	0.05
5	1	H	647	tight thermal	0.026	0.5
5	2	I	647	tight positional	0.025	0.05
5	2	I	647	tight thermal	0.032	0.5
5	3	J	647	tight positional	0.023	0.05
5	3	J	647	tight thermal	0.028	0.5
5	4	K	647	tight positional	0.02	0.05
5	4	K	647	tight thermal	0.026	0.5
5	5	L	647	tight positional	0.019	0.05
5	5	L	647	tight thermal	0.025	0.5
5	6	M	647	tight positional	0.024	0.05
5	6	M	647	tight thermal	0.032	0.5
5	7	N	647	tight positional	0.022	0.05
5	7	N	647	tight thermal	0.029	0.5
6	1	H	1398	tight positional	0.017	0.05
6	1	H	1398	tight thermal	0.014	0.5
6	2	I	1398	tight positional	0.017	0.05
6	2	I	1398	tight thermal	0.016	0.5
6	3	J	1398	tight positional	0.018	0.05
6	3	J	1398	tight thermal	0.017	0.5
6	4	K	1398	tight positional	0.028	0.05
6	4	K	1398	tight thermal	0.013	0.5
6	5	L	1398	tight positional	0.016	0.05
6	5	L	1398	tight thermal	0.014	0.5
6	6	M	1398	tight positional	0.028	0.05
6	6	M	1398	tight thermal	0.014	0.5
6	7	N	1398	tight positional	0.016	0.05
6	7	N	1398	tight thermal	0.014	0.5
7	1	O	728	tight positional	0.01	0.05
7	1	O	728	tight thermal	0.01	0.5
7	2	P	728	tight positional	0.01	0.05
7	2	P	728	tight thermal	0.01	0.5
7	3	Q	728	tight positional	0.01	0.05
7	3	Q	728	tight thermal	0.009	0.5
7	4	R	728	tight positional	0.01	0.05
7	4	R	728	tight thermal	0.01	0.5
7	5	S	728	tight positional	0.01	0.05
7	5	S	728	tight thermal	0.009	0.5
7	6	T	728	tight positional	0.01	0.05
7	6	T	728	tight thermal	0.01	0.5
7	7	U	728	tight positional	0.01	0.05
7	7	U	728	tight thermal	0.009	0.5

LS refinement shell

Resolution (Å)	R_factor R_free	Num. reflection R_free	R_factor R_work	Num. reflection R_work	Refine-ID
2.808-2.8807	0.433	219	0.385	11893	X-RAY DIFFRACTION
2.881-2.9594	0.38	287	0.365	13797	X-RAY DIFFRACTION
2.959-3.0448	0.388	311	0.329	15545	X-RAY DIFFRACTION
3.045-3.1382	0.341	360	0.324	17756	X-RAY DIFFRACTION
3.138-3.2407	0.317	362	0.311	18512	X-RAY DIFFRACTION
3.241-3.3539	0.341	387	0.307	18520	X-RAY DIFFRACTION
3.354-3.4799	0.304	352	0.299	18328	X-RAY DIFFRACTION
3.48-3.6212	0.265	367	0.269	17765	X-RAY DIFFRACTION
3.621-3.7813	0.308	351	0.291	17084	X-RAY DIFFRACTION
3.781-3.9647	0.281	340	0.279	16351	X-RAY DIFFRACTION
3.965-4.1777	0.229	303	0.229	15648	X-RAY DIFFRACTION
4.178-4.4291	0.24	284	0.217	14733	X-RAY DIFFRACTION
4.429-4.7322	0.251	285	0.214	13898	X-RAY DIFFRACTION
4.732-5.1075	0.23	258	0.219	12967	X-RAY DIFFRACTION
5.107-5.589	0.297	265	0.266	11932	X-RAY DIFFRACTION
5.589-6.2387	0.303	226	0.288	10823	X-RAY DIFFRACTION
6.239-7.1847	0.281	211	0.263	9576	X-RAY DIFFRACTION
7.185-8.7533	0.209	157	0.183	8161	X-RAY DIFFRACTION
8.753-12.189	0.136	121	0.151	6327	X-RAY DIFFRACTION
12.189-49.3865	0.263	71	0.271	3458	X-RAY DIFFRACTION

Refinement TLS params.

Method: refined / Refine-ID: X-RAY DIFFRACTION

ID	L11 (°²)	L12 (°²)	L13 (°²)	L22 (°²)	L23 (°²)	L33 (°²)	S11 (Å °)	S12 (Å °)	S13 (Å °)	S21 (Å °)	S22 (Å °)	S23 (Å °)	S31 (Å °)	S32 (Å °)	S33 (Å °)	T11 (Å²)	T12 (Å²)	T13 (Å²)	T22 (Å²)	T23 (Å²)	T33 (Å²)	Origin x (Å)	Origin y (Å)	Origin z (Å)
1	5.8885	-0.3703	0.1609	3.7299	-0.3795	0.4263	-0.0461	0.1348	-0.0621	0.0526	0.0645	-0.2	0.0378	-0.1339	-0.0184	0.6433	-0.0205	0.0047	0.655	-0.0658	0.6271	33.185	-53.269	-12.439
2	11.8107	7.2608	0.7235	7.7613	-0.5418	2.8378	-0.5361	1.0202	-0.0576	-0.8661	0.5731	-0.5001	-0.372	-0.072	-0.037	0.8519	0.0661	0.0587	0.9091	-0.0361	1.298	30.136	-34.343	-31.393
3	9.4327	1.2683	-4.4337	6.7806	1.0627	24.86	0.0137	0.0256	-0.0139	-0.0821	-0.1701	0.0005	0.6277	1.0312	0.1564	2.0964	-0.2807	0.248	2.0387	-0.4299	1.8276	37.643	-37.18	-62.092
4	5.5628	-0.2837	0.5278	3.3632	-0.1272	0.6819	-0.0547	-0.0294	-0.107	-0.0207	-0.0125	0.158	0.1491	-0.1017	0.0672	0.6501	-0.0255	0.0547	0.647	-0.1052	1.0087	51.401	-88.96	-12.602
5	2.3641	-2.389	0.1715	12.2987	-0.2187	3.3299	0.0577	0.7401	0.1478	-1.1558	-0.1917	0.0677	-0.1934	-0.3678	0.134	0.8378	-0.0151	0.062	1.1117	-0.0985	1.6293	34.656	-79.515	-31.398
6	2.7851	1.9535	-4.3115	7.7728	-5.7554	26.1696	-0.1448	-0.1674	-0.3395	0.0689	-0.1533	0.0259	0.4273	1.6507	0.2981	2.0182	0.1841	0.4267	2.2746	-0.1603	2.1062	41.483	-75.323	-62.156
7	4.5444	1.3104	0.1649	4.4307	0.4353	0.3508	0.0411	-0.0344	0.018	-0.0048	-0.0236	0.1248	0.1047	0.0357	-0.0176	0.6453	0.0121	0.0569	0.6419	0.0035	0.7697	90.794	-96.793	-12.646
8	8.1987	-3.9356	0.2239	8.372	0.1441	4.8157	0.221	0.9336	0.1622	-1	-0.2681	0.1846	-0.0619	-0.513	0.0471	0.9042	-0.0706	-0.0063	0.8914	-0.0073	1.4752	72.9235	-104.0459	-31.4786
9	6.6836	-1.9234	2.4125	6.6652	-5.8917	23.7504	-0.1034	0.0745	-0.1847	0.4606	0.1992	0.27	-1.5914	0.1165	-0.0957	2.0485	0.1179	0.4154	2.2185	0.1671	1.8948	74.091	-95.762	-62.034
10	4.3233	-0.1511	-0.0062	4.3328	0.1298	0.4015	-0.0098	0.1205	0.126	0.0894	-0.0253	0.3376	0.0151	0.1086	0.0351	0.6139	-0.0057	-0.0014	0.7038	0.0745	0.7662	121.473	-70.925	-12.322
11	13.0139	2.0565	-0.5735	4.0501	0.4725	3.4508	-0.2293	1.1708	0.6102	-0.5414	0.1411	0.7963	0.3193	0.0094	0.0882	0.8757	0.0154	-0.0173	0.8866	0.0712	1.3767	115.984	-89.352	-31.206
12	8.8212	0.7758	5.7725	5.0146	-1.6317	25.3982	-0.0172	-0.2538	-0.2512	-0.246	-0.3016	-0.23	-0.7975	-0.7072	0.3189	1.9962	-0.2683	0.0388	2.214	0.5595	1.8973	110.465	-83.365	-61.965
13	5.154	-1.0539	-0.4339	4.0143	0.2474	0.6684	-0.0663	0.0591	0.3265	-0.0133	0.0088	-0.1931	-0.1364	0.1515	0.0575	0.6318	-0.0435	-0.0558	0.7076	0.1586	1.4438	120.61	-30.801	-12.506
14	2.8528	4.8506	-0.9257	11.1249	0.4258	4.3049	-0.2177	0.8506	-0.0765	-1.1491	0.2114	-0.0877	0.2453	0.0674	0.0063	0.8713	0.089	-0.0553	1.1138	0.1818	1.8437	131.571	-46.663	-31.227
15	6.4452	2.0538	5.8214	10.7446	4.1698	21.5568	-0.2553	0.0279	0.4355	0.5275	-0.0068	-0.3868	-0.7731	-0.9213	0.2621	2.1521	0.0627	-0.2302	2.2789	0.4906	1.9161	123.434	-47.191	-61.965
16	4.6385	0.8009	-0.2603	3.3099	0.0346	0.6757	-0.0616	0.0523	0.3493	-0.0353	0.0039	-0.3346	-0.2047	0.0482	0.0577	0.7388	-0.0009	-0.1135	0.6504	0.1138	1.4913	88.628	-6.513	-12.689
17	7.154	-7.7232	-0.1676	12.1429	-1.3652	3.4842	0.472	1.0822	-0.2766	-1.2111	-0.433	-0.0462	0.3678	0.3727	-0.039	0.9716	-0.2572	0.0099	1.2315	0.1539	2.2423	107.982	-7.676	-31.421
18	4.9129	-2.25	1.7619	7.2552	3.6195	19.0307	0.0936	-0.1704	0.5237	0.8461	0.085	-0.5889	1.1263	-0.6084	-0.1786	2.3676	0.0767	-0.3411	2.4637	0.1042	2.3336	103.431	-14.18	-62.171
19	2.8629	0.4556	-0.0091	5.3457	-0.3907	0.6166	0.0316	-0.0252	0.1373	0.024	0.0177	-0.1212	-0.0934	-0.0084	-0.0492	0.6447	0.021	-0.063	0.6313	-0.0094	0.9284	49.609	-16.648	-12.852
20	12.95	-1.6757	-1.6902	4.8089	-0.5785	4.6782	0.1571	0.9865	-0.424	-0.8359	-0.1642	-0.733	-0.0994	0.4161	0.0071	1.034	-0.0157	0.0534	0.792	-0.0311	1.6641	62.599	-2.379	-31.713
21	9.3518	0.2348	-4.6647	4.0039	4.9173	27.5855	-0.1294	-0.31	0.548	0.3059	0.0556	0.074	1.4703	0.5305	0.0738	2.0562	-0.1049	-0.0175	2.1241	-0.2363	2.1269	65.199	-10.009	-62.31
22	3.8783	0.4356	0.051	6.0879	0.179	1.2499	-0.0791	0.003	-0.4887	-0.0974	0.0694	0.3223	0.1171	-0.213	0.0097	0.6488	-0.0104	0.0705	0.7088	0.0314	0.8872	40.548	-76.5	19.525
23	8.0247	-3.4066	7.4544	3.9366	-2.7032	9.819	0.4246	-0.1377	0.0061	0.3733	-0.3229	-0.1195	0.8067	0.4977	-0.1017	1.3668	-0.0996	0.1646	1.2589	0.0709	1.5481	38.316	-93.283	44.73
24	8.5657	1.9623	-0.4832	9.473	0.4526	5.3539	-0.1363	-0.1433	0.7625	0.4333	-0.021	-0.3565	-0.4251	0.0194	0.1573	1.5869	0.0238	-0.0763	1.4817	-0.029	1.8059	40.556	-70.595	59.281
25	4.4373	-0.9665	-0.2742	5.8531	0.4882	0.6696	0.1165	0.1161	-0.0499	-0.0186	-0.0869	-0.0924	0.0708	-0.0803	-0.0296	0.618	0.0023	0.0049	0.6719	0.026	0.7351	74.158	-97.46	19.523
26	6.9721	0.4346	3.5521	0.1445	3.7757	10.0901	-0.1331	-0.4843	-0.0336	0.6898	0.2571	-0.1885	0.2181	0.9033	-0.124	1.2043	0.0179	-0.0527	1.1067	0.1457	1.54	85.969	-109.9817	44.7359
27	7.9965	-0.6232	0.1941	9.7848	0.1546	5.5883	-0.358	-0.9293	0.896	0.579	0.4305	0.4484	-0.2439	-0.3675	-0.0725	1.4159	0.1205	0.0083	1.6128	-0.0193	1.6768	69.386	-94.681	59.318
28	8.1352	0.1122	-0.1944	4.1457	0.3057	0.647	-0.1674	0.1962	-0.2678	0.0553	0.1624	-0.0634	0.0831	0.0925	0.0049	0.6452	-0.0109	0.0073	0.7334	0.0072	0.7159	111.762	-84.492	19.723
29	4.4728	-1.0638	2.0026	7.0482	5.3818	9.426	-0.2908	-0.8429	-0.0557	-0.0595	0.018	-0.1484	-0.9508	0.4673	0.2729	1.0471	-0.0078	0.0084	1.4368	0.139	1.493	128.165	-83.062	45.14
30	11.6188	-0.8895	1.8511	6.9637	-1.3512	6.803	0.5314	-1.5075	0.4072	0.1929	-0.6813	1.1678	0.0299	-0.3522	0.1499	1.2321	-0.1504	0.2434	1.8844	-0.2208	1.7875	105.864	-86.255	59.848
31	5.0455	1.5001	-0.6393	5.4894	-0.6023	1.3217	-0.087	-0.0503	0.1569	-0.0471	-0.0602	-0.5318	-0.0837	0.2132	0.1471	0.6507	-0.0088	-0.1082	0.7187	0.0134	1.2699	124.883	-46.593	19.63
32	4.9815	-2.1763	-3.5785	1.8153	3.7529	10.448	0.562	-0.5659	0.397	0.0214	-0.3342	-0.3567	-1.3324	-0.0573	-0.2278	1.4555	-0.2973	-0.3195	1.3788	0.019	2.0505	133.697	-32.734	44.875
33	8.5805	0.0354	1.2215	7.9596	-0.7227	6.9737	-0.0412	-0.1533	-0.4944	0.8714	-0.3345	1.0044	0.2469	-0.114	0.3757	1.5945	-0.1802	0.0735	1.5014	-0.0792	2.0872	121.949	-52.203	59.46
34	4.7116	-1.7527	0.4527	7.2596	-0.9948	1.7065	0.0794	0.1764	0.6287	-0.0645	-0.2415	-0.5814	-0.3648	0.1736	0.1621	0.7133	-0.0195	-0.1223	0.7095	-0.0529	1.4899	103.652	-12.846	19.735
35	1.22	-1.3265	-4.2233	1.0639	1.2898	8.8085	-0.1988	-0.1098	0.2752	0.975	0.06	-0.0338	-0.6044	-0.698	0.1388	1.9188	-0.1058	-0.3133	1.3319	-0.1158	2.1877	97.821	3.141	44.837
36	7.0358	0.6209	-0.4433	8.2168	0.4467	5.7255	-0.5805	-0.3131	-0.584	0.6147	0.412	-0.0995	-0.0485	0.123	0.1685	2.0415	0.0735	-0.2108	1.5102	-0.0501	2.1085	105.642	-18.357	59.337
37	9.8555	-1.9496	0.5839	10.279	-0.818	0.0046	0.0002	0.4249	-0.2259	0.2713	-0.0087	-0.271	-0.1355	0.0541	0.0085	0.7393	-0.007	-0.0439	0.7938	-0.0561	0.0066	64.066	-8.702	19.43
38	2.8564	5.1233	-1.306	7.4215	-6.0653	9.4077	-0.1136	-0.6417	-0.1377	1.1545	0.294	0.136	0.1584	-0.9532	-0.1804	1.5177	0.0114	0.0878	1.2041	-0.0238	1.4916	47.644	-2.762	44.567
39	8.512	-1.7099	0.8062	7.7958	0.2076	7.0016	-0.0269	-1.1102	-0.6355	0.2983	-0.0361	-0.8351	-0.0699	0.4445	0.063	1.7406	-0.0966	-0.0786	1.8085	0.0393	1.716	69.255	-9.246	59.224
40	8.5881	1.8821	0.0255	5.6588	0.0601	0.3428	-0.3011	0.0876	0.477	-0.1062	0.2898	0.3227	-0.0555	-0.0841	0.0114	0.7072	0	-0.0473	0.7062	-0.0073	0.6715	35.915	-36.784	19.642
41	2.9445	-0.1691	-0.8402	7.5564	-5.1869	10.0945	0.2561	-0.8228	-0.0759	0.1413	-0.0506	-0.2688	0.959	0.0414	-0.2055	1.217	-0.0303	-0.0428	1.4887	-0.2608	1.4137	21.694	-45.429	45.027
42	9.2506	0.1644	-2.1397	8.7002	1.077	7.1683	0.9783	-0.9955	-0.3545	0.9749	-1.0963	-1.3073	-0.5106	0.3867	0.118	1.704	-0.3713	-0.3773	1.7652	0.1435	1.7552	40.326	-32.887	59.705
43	15.5516	3.0717	-8.2353	7.3133	-2.6989	13.0437	0.621	-0.1299	0.2218	0.9347	-0.2677	0.5229	-0.8462	-0.3105	-0.3533	2.1824	-0.045	-0.0565	2.1042	-0.035	1.7581	65.633	-29.496	-98.12
44	6.97	4.7764	-3.0862	13.545	-8.4018	10.726	0.0338	-0.383	-0.3875	0.8719	0.109	0.2751	-0.3128	-0.2475	-0.1428	2.0615	0.0105	0.0192	2.4056	-0.2492	1.7144	52.952	-48.79	-98.054
45	8.5979	-4.4944	4.4573	11.0979	-6.7275	13.4846	-0.1342	-0.8968	-0.7881	0.3078	0.1512	0.0894	0.078	-0.3097	-0.0171	2.0705	-0.1505	0.1783	2.2338	-0.1035	1.864	60.224	-70.818	-98.118
46	13.5842	-0.8156	5.8808	3.7277	-1.0751	12.5549	0.4314	-0.8084	-0.2715	0.2947	-0.268	-0.5858	0.3405	-0.0751	-0.1634	2.2769	0.0038	0.1222	2.1737	0.0888	1.8347	81.924	-78.475	-98.043
47	15.3907	5.4344	8.7759	10.1961	6.0207	13.3935	0.2786	-0.3969	0.1266	0.8007	0.0275	-0.6567	0.7571	0.2201	-0.306	2.1137	0.1675	0.2357	2.2191	0.2832	1.7912	101.601	-66.509	-97.982
48	5.9086	-1.5104	-0.4552	18.4496	8.0374	14.4428	0.0025	-0.5372	0.8741	1.4887	0.4359	-0.9326	0.764	0.6393	-0.4384	1.9386	-0.0195	0.0225	2.3411	0.265	1.9187	104.5	-43.639	-98.161
49	11.2186	-2.7902	-4.5813	6.2603	4.6242	11.2222	0.0157	-1.3565	0.5616	0.7325	0.1281	-0.2445				2.0358	-0.1535	-0.046	2.2887	0.1263	1.9506	88.527	-26.924	-98.327

Refinement TLS group

ID	Refine-ID	Refine TLS-ID	Auth asym-ID	Label asym-ID	Auth seq-ID	Label seq-ID
1	X-RAY DIFFRACTION	1	A	A	2 - 135	1 - 134
2	X-RAY DIFFRACTION	1	A	A	410 - 525	409 - 524
3	X-RAY DIFFRACTION	2	A	A	136 - 190	135 - 189
4	X-RAY DIFFRACTION	2	A	A	375 - 409	374 - 408
5	X-RAY DIFFRACTION	3	A	A	191 - 374	190 - 373
6	X-RAY DIFFRACTION	4	B	B	2 - 135	1 - 134
7	X-RAY DIFFRACTION	4	B	B	410 - 525	409 - 524
8	X-RAY DIFFRACTION	5	B	B	136 - 190	135 - 189
9	X-RAY DIFFRACTION	5	B	B	375 - 409	374 - 408
10	X-RAY DIFFRACTION	6	B	B	191 - 374	190 - 373
11	X-RAY DIFFRACTION	7	C	C	2 - 135	1 - 134
12	X-RAY DIFFRACTION	7	C	C	410 - 525	409 - 524
13	X-RAY DIFFRACTION	8	C	C	136 - 190	135 - 189
14	X-RAY DIFFRACTION	8	C	C	375 - 409	374 - 408
15	X-RAY DIFFRACTION	9	C	C	191 - 374	190 - 373
16	X-RAY DIFFRACTION	10	D	D	2 - 135	1 - 134
17	X-RAY DIFFRACTION	10	D	D	410 - 525	409 - 524
18	X-RAY DIFFRACTION	11	D	D	136 - 190	135 - 189
19	X-RAY DIFFRACTION	11	D	D	375 - 409	374 - 408
20	X-RAY DIFFRACTION	12	D	D	191 - 374	190 - 373
21	X-RAY DIFFRACTION	13	E	E	2 - 135	1 - 134
22	X-RAY DIFFRACTION	13	E	E	410 - 525	409 - 524
23	X-RAY DIFFRACTION	14	E	E	136 - 190	135 - 189
24	X-RAY DIFFRACTION	14	E	E	375 - 409	374 - 408
25	X-RAY DIFFRACTION	15	E	E	191 - 374	190 - 373
26	X-RAY DIFFRACTION	16	F	F	2 - 135	1 - 134
27	X-RAY DIFFRACTION	16	F	F	410 - 525	409 - 524
28	X-RAY DIFFRACTION	17	F	F	136 - 190	135 - 189
29	X-RAY DIFFRACTION	17	F	F	375 - 409	374 - 408
30	X-RAY DIFFRACTION	18	F	F	191 - 374	190 - 373
31	X-RAY DIFFRACTION	19	G	G	2 - 135	1 - 134
32	X-RAY DIFFRACTION	19	G	G	410 - 525	409 - 524
33	X-RAY DIFFRACTION	20	G	G	136 - 190	135 - 189
34	X-RAY DIFFRACTION	20	G	G	375 - 409	374 - 408
35	X-RAY DIFFRACTION	21	G	G	191 - 374	190 - 373
36	X-RAY DIFFRACTION	22	H	H	2 - 135	1 - 134
37	X-RAY DIFFRACTION	22	H	H	410 - 525	409 - 524
38	X-RAY DIFFRACTION	23	H	H	136 - 190	135 - 189
39	X-RAY DIFFRACTION	23	H	H	375 - 409	374 - 408
40	X-RAY DIFFRACTION	24	H	H	191 - 374	190 - 373
41	X-RAY DIFFRACTION	25	I	I	2 - 135	1 - 134
42	X-RAY DIFFRACTION	25	I	I	410 - 525	409 - 524
43	X-RAY DIFFRACTION	26	I	I	136 - 190	135 - 189
44	X-RAY DIFFRACTION	26	I	I	375 - 409	374 - 408
45	X-RAY DIFFRACTION	27	I	I	191 - 374	190 - 373
46	X-RAY DIFFRACTION	28	J	J	2 - 135	1 - 134
47	X-RAY DIFFRACTION	28	J	J	410 - 525	409 - 524
48	X-RAY DIFFRACTION	29	J	J	136 - 190	135 - 189
49	X-RAY DIFFRACTION	29	J	J	375 - 409	374 - 408
50	X-RAY DIFFRACTION	30	J	J	191 - 374	190 - 373
51	X-RAY DIFFRACTION	31	K	K	2 - 135	1 - 134
52	X-RAY DIFFRACTION	31	K	K	410 - 525	409 - 524
53	X-RAY DIFFRACTION	32	K	K	136 - 190	135 - 189
54	X-RAY DIFFRACTION	32	K	K	375 - 409	374 - 408
55	X-RAY DIFFRACTION	33	K	K	191 - 374	190 - 373
56	X-RAY DIFFRACTION	34	L	L	2 - 135	1 - 134
57	X-RAY DIFFRACTION	34	L	L	410 - 525	409 - 524
58	X-RAY DIFFRACTION	35	L	L	136 - 190	135 - 189
59	X-RAY DIFFRACTION	35	L	L	375 - 409	374 - 408
60	X-RAY DIFFRACTION	36	L	L	191 - 374	190 - 373
61	X-RAY DIFFRACTION	37	M	M	2 - 135	1 - 134
62	X-RAY DIFFRACTION	37	M	M	410 - 525	409 - 524
63	X-RAY DIFFRACTION	38	M	M	136 - 190	135 - 189
64	X-RAY DIFFRACTION	38	M	M	375 - 409	374 - 408
65	X-RAY DIFFRACTION	39	M	M	191 - 374	190 - 373
66	X-RAY DIFFRACTION	40	N	N	2 - 135	1 - 134
67	X-RAY DIFFRACTION	40	N	N	410 - 525	409 - 524
68	X-RAY DIFFRACTION	41	N	N	136 - 190	135 - 189
69	X-RAY DIFFRACTION	41	N	N	375 - 409	374 - 408
70	X-RAY DIFFRACTION	42	N	N	191 - 374	190 - 373
71	X-RAY DIFFRACTION	43	O	O	1 - 97	1 - 97
72	X-RAY DIFFRACTION	44	P	P	1 - 97	1 - 97
73	X-RAY DIFFRACTION	45	Q	Q	1 - 97	1 - 97
74	X-RAY DIFFRACTION	46	R	R	1 - 97	1 - 97
75	X-RAY DIFFRACTION	47	S	S	1 - 97	1 - 97
76	X-RAY DIFFRACTION	48	T	T	1 - 97	1 - 97
77	X-RAY DIFFRACTION	49	U	U	1 - 97	1 - 97

Software

*PLUS

Name: REFMAC / Version: 5.1.24 / Classification: refinement

Refinement

*PLUS

Highest resolution: 2.8 Å / Lowest resolution: 50 Å / % reflection R_free: 2 % / R_factor R_free: 0.278 / R_factor R_work: 0.262

Solvent computation

*PLUS

Displacement parameters

*PLUS

Refine LS restraints

*PLUS

Refine-ID	Type	Dev ideal
X-RAY DIFFRACTION	p_bond_d	0.013
X-RAY DIFFRACTION	p_angle_d
X-RAY DIFFRACTION	p_angle_deg	1.362

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