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- PDB-5j9q: Crystal structure of the NuA4 core complex -

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Basic information

Entry
Database: PDB / ID: 5j9q
TitleCrystal structure of the NuA4 core complex
Components
  • Chromatin modification-related protein EAF6
  • Chromatin modification-related protein YNG2
  • Enhancer of polycomb-like protein 1
  • Histone acetyltransferase ESA1
  • Htz1
KeywordsTRANSFERASE / NuA4 / nucleosome / histone / acetylation
Function / homology
Function and homology information


PI5P Regulates TP53 Acetylation / NuA3b histone acetyltransferase complex / NuA3a histone acetyltransferase complex / NuA3 histone acetyltransferase complex / DNA Damage/Telomere Stress Induced Senescence / Sensing of DNA Double Strand Breaks / piccolo histone acetyltransferase complex / peptide 2-hydroxyisobutyryltransferase activity / histone crotonyltransferase activity / nuclear-transcribed mRNA catabolic process, non-stop decay ...PI5P Regulates TP53 Acetylation / NuA3b histone acetyltransferase complex / NuA3a histone acetyltransferase complex / NuA3 histone acetyltransferase complex / DNA Damage/Telomere Stress Induced Senescence / Sensing of DNA Double Strand Breaks / piccolo histone acetyltransferase complex / peptide 2-hydroxyisobutyryltransferase activity / histone crotonyltransferase activity / nuclear-transcribed mRNA catabolic process, non-stop decay / positive regulation of triglyceride biosynthetic process / Condensation of Prophase Chromosomes / SIRT1 negatively regulates rRNA expression / SUMOylation of transcription cofactors / Activated PKN1 stimulates transcription of AR (androgen receptor) regulated genes KLK2 and KLK3 / DNA-templated transcription elongation / Assembly of the ORC complex at the origin of replication / histone H4 acetyltransferase activity / rDNA heterochromatin formation / Recruitment and ATM-mediated phosphorylation of repair and signaling proteins at DNA double strand breaks / Oxidative Stress Induced Senescence / histone H3K4me3 reader activity / silent mating-type cassette heterochromatin formation / protein-lysine-acetyltransferase activity / RNA Polymerase I Promoter Escape / NuA4 histone acetyltransferase complex / : / Estrogen-dependent gene expression / positive regulation of macroautophagy / histone acetyltransferase activity / histone acetyltransferase / Transferases; Acyltransferases; Transferring groups other than aminoacyl groups / meiotic cell cycle / transcription coregulator activity / transcription elongation by RNA polymerase II / positive regulation of transcription elongation by RNA polymerase II / euchromatin / chromatin DNA binding / structural constituent of chromatin / nucleosome / heterochromatin formation / chromatin organization / regulation of cell cycle / RNA polymerase II cis-regulatory region sequence-specific DNA binding / chromatin remodeling / protein heterodimerization activity / DNA repair / DNA-templated transcription / chromatin binding / regulation of DNA-templated transcription / regulation of transcription by RNA polymerase II / chromatin / zinc ion binding / identical protein binding / nucleus / cytosol
Similarity search - Function
N-acetyl transferase-like / Chromatin modification-related protein Eaf6 / Histone acetyltransferase subunit NuA4 / Inhibitor of growth protein, N-terminal histone-binding / Inhibitor of growth proteins N-terminal histone-binding / Inhibitor of growth proteins N-terminal histone-binding / ING family / Enhancer of polycomb protein / : / MYST, zinc finger domain ...N-acetyl transferase-like / Chromatin modification-related protein Eaf6 / Histone acetyltransferase subunit NuA4 / Inhibitor of growth protein, N-terminal histone-binding / Inhibitor of growth proteins N-terminal histone-binding / Inhibitor of growth proteins N-terminal histone-binding / ING family / Enhancer of polycomb protein / : / MYST, zinc finger domain / MYST family zinc finger domain / Histone acetyltransferase domain, MYST-type / MOZ/SAS family / MYST-type histone acetyltransferase (HAT) domain profile. / RNA binding activity-knot of a chromodomain / RNA binding activity-knot of a chromodomain / Wheat Germ Agglutinin (Isolectin 2); domain 1 / Chromo/chromo shadow domain / Chromatin organization modifier domain / Enhancer of polycomb-like, N-terminal / Enhancer of polycomb-like / Chromo-like domain superfamily / Gcn5-related N-acetyltransferase (GNAT) / Zinc finger, PHD-type, conserved site / PHD-finger / Acyl-CoA N-acyltransferase / Zinc finger PHD-type signature. / Aminopeptidase / Zinc finger PHD-type profile. / Zinc finger, PHD-finger / Zinc finger, PHD-type / PHD zinc finger / Histone H2A conserved site / Histone H2A signature. / Histone H2A, C-terminal domain / C-terminus of histone H2A / Histone 2A / Histone H2A / Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain / Zinc finger, FYVE/PHD-type / Histone H2A/H2B/H3 / Core histone H2A/H2B/H3/H4 domain / Histone-fold / Arc Repressor Mutant, subunit A / Zinc finger, RING/FYVE/PHD-type / Winged helix-like DNA-binding domain superfamily / 2-Layer Sandwich / Orthogonal Bundle / 3-Layer(aba) Sandwich / Mainly Alpha / Alpha Beta
Similarity search - Domain/homology
Chromatin modification-related protein YNG2 / Enhancer of polycomb-like protein 1 / Chromatin modification-related protein EAF6 / Histone acetyltransferase ESA1 / Histone H2A.Z
Similarity search - Component
Biological speciesSaccharomyces cerevisiae (brewer's yeast)
MethodX-RAY DIFFRACTION / SYNCHROTRON / MOLECULAR REPLACEMENT / Resolution: 3.25 Å
AuthorsChen, Z.C. / Xu, P.
Funding support China, 1items
OrganizationGrant numberCountry
Ministry of Science and Technology (China)2014CB910100 China
CitationJournal: Mol Cell / Year: 2016
Title: The NuA4 Core Complex Acetylates Nucleosomal Histone H4 through a Double Recognition Mechanism.
Authors: Peng Xu / Chengmin Li / Zhihong Chen / Shuanying Jiang / Shilong Fan / Jiawei Wang / Junbiao Dai / Ping Zhu / Zhucheng Chen /
Abstract: NuA4 catalyzes the acetylation of nucleosomes at histone H4, which is a well-established epigenetic event, controlling many genomic processes in Saccharomyces cerevisiae. Here we report the crystal ...NuA4 catalyzes the acetylation of nucleosomes at histone H4, which is a well-established epigenetic event, controlling many genomic processes in Saccharomyces cerevisiae. Here we report the crystal structures of the NuA4 core complex and a cryoelectron microscopy structure with the nucleosome. The structures show that the histone-binding pocket of the enzyme is rearranged, suggesting its activation. The enzyme binds the histone tail mainly through the target lysine residue, with a preference for a small residue at the -1 position. The complex engages the nucleosome at the dish face and orients its catalytic pocket close to the H4 tail to achieve selective acetylation. The combined data reveal a space-sequence double recognition mechanism of the histone tails by a modifying enzyme in the context of the nucleosome.
History
DepositionApr 11, 2016Deposition site: RCSB / Processing site: PDBJ
Revision 1.0Oct 26, 2016Provider: repository / Type: Initial release
Revision 1.1Oct 18, 2017Group: Author supporting evidence / Database references / Derived calculations
Category: citation / pdbx_audit_support / pdbx_struct_oper_list
Item: _citation.journal_id_CSD / _pdbx_audit_support.funding_organization / _pdbx_struct_oper_list.symmetry_operation
Revision 1.2Oct 30, 2024Group: Data collection / Database references / Structure summary
Category: chem_comp_atom / chem_comp_bond ...chem_comp_atom / chem_comp_bond / database_2 / pdbx_entry_details / pdbx_modification_feature
Item: _database_2.pdbx_DOI / _database_2.pdbx_database_accession

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Structure visualization

Structure viewerMolecule:
MolmilJmol/JSmol

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Assembly

Deposited unit
E: Histone acetyltransferase ESA1
F: Chromatin modification-related protein EAF6
G: Enhancer of polycomb-like protein 1
A: Histone acetyltransferase ESA1
I: Histone acetyltransferase ESA1
N: Enhancer of polycomb-like protein 1
C: Enhancer of polycomb-like protein 1
D: Chromatin modification-related protein YNG2
B: Chromatin modification-related protein EAF6
J: Chromatin modification-related protein EAF6
H: Chromatin modification-related protein YNG2
K: Chromatin modification-related protein YNG2
L: Htz1
M: Htz1
O: Htz1


Theoretical massNumber of molelcules
Total (without water)318,34715
Polymers318,34715
Non-polymers00
Water00
1
E: Histone acetyltransferase ESA1
F: Chromatin modification-related protein EAF6
G: Enhancer of polycomb-like protein 1
H: Chromatin modification-related protein YNG2
L: Htz1


Theoretical massNumber of molelcules
Total (without water)106,1165
Polymers106,1165
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area19400 Å2
ΔGint-108 kcal/mol
Surface area41400 Å2
MethodPISA
2
A: Histone acetyltransferase ESA1
C: Enhancer of polycomb-like protein 1
D: Chromatin modification-related protein YNG2
B: Chromatin modification-related protein EAF6
O: Htz1


Theoretical massNumber of molelcules
Total (without water)106,1165
Polymers106,1165
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area18780 Å2
ΔGint-108 kcal/mol
Surface area41640 Å2
MethodPISA
3
I: Histone acetyltransferase ESA1
N: Enhancer of polycomb-like protein 1
J: Chromatin modification-related protein EAF6
K: Chromatin modification-related protein YNG2
M: Htz1


Theoretical massNumber of molelcules
Total (without water)106,1165
Polymers106,1165
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area18750 Å2
ΔGint-111 kcal/mol
Surface area41000 Å2
MethodPISA
Unit cell
Length a, b, c (Å)217.857, 190.990, 172.346
Angle α, β, γ (deg.)90.00, 113.56, 90.00
Int Tables number5
Space group name H-MC121

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Components

#1: Protein Histone acetyltransferase ESA1


Mass: 36556.883 Da / Num. of mol.: 3 / Fragment: UNP residues 141-445 / Mutation: E338Q
Source method: isolated from a genetically manipulated source
Source: (gene. exp.) Saccharomyces cerevisiae (strain ATCC 204508 / S288c) (yeast)
Strain: ATCC 204508 / S288c / Gene: ESA1, YOR244W, O5257 / Production host: Escherichia coli (E. coli) / References: UniProt: Q08649, histone acetyltransferase
#2: Protein Chromatin modification-related protein EAF6 / ESA1-associated factor 6


Mass: 12915.704 Da / Num. of mol.: 3
Source method: isolated from a genetically manipulated source
Source: (gene. exp.) Saccharomyces cerevisiae (strain ATCC 204508 / S288c) (yeast)
Strain: ATCC 204508 / S288c / Gene: EAF6, YJR082C, J1854 / Production host: Escherichia coli (E. coli) / References: UniProt: P47128
#3: Protein Enhancer of polycomb-like protein 1


Mass: 41835.215 Da / Num. of mol.: 3 / Fragment: UNP residues 50-400
Source method: isolated from a genetically manipulated source
Source: (gene. exp.) Saccharomyces cerevisiae (strain ATCC 204508 / S288c) (yeast)
Strain: ATCC 204508 / S288c / Gene: EPL1, YFL024C / Production host: Escherichia coli (E. coli) / References: UniProt: P43572
#4: Protein Chromatin modification-related protein YNG2 / ESA1-associated factor 4 / ING1 homolog 2


Mass: 13828.894 Da / Num. of mol.: 3 / Fragment: UNP residues 1-120
Source method: isolated from a genetically manipulated source
Source: (gene. exp.) Saccharomyces cerevisiae (strain ATCC 204508 / S288c) (yeast)
Strain: ATCC 204508 / S288c / Gene: YNG2, EAF4, NBN1, YHR090C / Production host: Escherichia coli (E. coli) / References: UniProt: P38806
#5: Protein/peptide Htz1


Mass: 979.049 Da / Num. of mol.: 3
Source method: isolated from a genetically manipulated source
Source: (gene. exp.) Saccharomyces cerevisiae (brewer's yeast)
Production host: Escherichia coli (E. coli) / References: UniProt: Q12692*PLUS
Has protein modificationY

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Experimental details

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Experiment

ExperimentMethod: X-RAY DIFFRACTION / Number of used crystals: 1

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Sample preparation

CrystalDensity Matthews: 5.16 Å3/Da / Density % sol: 76.17 %
Crystal growTemperature: 289 K / Method: evaporation
Details: 100mM NaCitrate (pH 6.5), 1.79M ammonium sulfate, 5%(w/v) glycerol

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Data collection

DiffractionMean temperature: 173 K
Diffraction sourceSource: SYNCHROTRON / Site: SSRF / Beamline: BL17U / Wavelength: 0.987 Å
DetectorType: ADSC QUANTUM 4r / Detector: CCD / Date: Jan 10, 2014
RadiationProtocol: MAD / Monochromatic (M) / Laue (L): M / Scattering type: x-ray
Radiation wavelengthWavelength: 0.987 Å / Relative weight: 1
ReflectionResolution: 3.25→30 Å / Num. obs: 79960 / % possible obs: 79.6 % / Redundancy: 3.4 % / Net I/σ(I): 6.9

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Processing

Software
NameVersionClassification
PHENIX(1.10.1_2155: ???)refinement
HKL-2000data reduction
HKL-2000data scaling
autoSHARPphasing
RefinementMethod to determine structure: MOLECULAR REPLACEMENT / Resolution: 3.25→29.996 Å / SU ML: 0.45 / Cross valid method: FREE R-VALUE / σ(F): 1.45 / Phase error: 30.12
RfactorNum. reflection% reflection
Rfree0.2731 1998 2.5 %
Rwork0.2502 --
obs0.2508 79960 78.86 %
Solvent computationShrinkage radii: 0.9 Å / VDW probe radii: 1.11 Å
Refinement stepCycle: LAST / Resolution: 3.25→29.996 Å
ProteinNucleic acidLigandSolventTotal
Num. atoms19671 0 0 0 19671
Refine LS restraints
Refine-IDTypeDev idealNumber
X-RAY DIFFRACTIONf_bond_d0.00420081
X-RAY DIFFRACTIONf_angle_d0.62127039
X-RAY DIFFRACTIONf_dihedral_angle_d14.1512277
X-RAY DIFFRACTIONf_chiral_restr0.0422898
X-RAY DIFFRACTIONf_plane_restr0.0033485
LS refinement shell
Resolution (Å)Rfactor RfreeNum. reflection RfreeRfactor RworkNum. reflection RworkRefine-ID% reflection obs (%)
3.2501-3.33130.3934510.32091972X-RAY DIFFRACTION28
3.3313-3.42120.3551800.33243164X-RAY DIFFRACTION45
3.4212-3.52180.36821060.31974093X-RAY DIFFRACTION58
3.5218-3.63530.3261240.30844876X-RAY DIFFRACTION70
3.6353-3.76490.33961460.29955710X-RAY DIFFRACTION81
3.7649-3.91540.29481590.28846187X-RAY DIFFRACTION87
3.9154-4.09320.30531600.26946231X-RAY DIFFRACTION89
4.0932-4.30840.27371640.26616399X-RAY DIFFRACTION91
4.3084-4.57740.23611640.25116405X-RAY DIFFRACTION91
4.5774-4.92940.2661650.23346446X-RAY DIFFRACTION91
4.9294-5.42280.26681700.23256626X-RAY DIFFRACTION94
5.4228-6.20150.30831720.24636723X-RAY DIFFRACTION95
6.2015-7.79050.25321710.23526664X-RAY DIFFRACTION94
7.7905-29.99750.19361660.18146466X-RAY DIFFRACTION90
Refinement TLS params.

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

IDL11 (°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)
10.0064-0.01940.00960.0140.00210.03810.05550.1545-0.0225-0.06730.0365-0.03140.02710.0303-00.2489-0.01410.03990.1294-0.05960.351310.830659.186875.444
20.0321-0.02850.00950.0128-0.00510.0026-0.05180.06980.0278-0.08450.0559-0.10670.17280.108400.0225-0.36320.2474-0.28240.05640.345911.545774.144670.5095
30.0031-0.00620.00060.01450.00520.01240.0902-0.11650.02490.01520.00980.03860.18620.18760-0.5588-0.44890.4632-0.26950.3826-0.22785.505173.468584.0465
40.00190.00140.00160.001400.0013-0.004-0.0202-0.0065-0.0157-0.0322-0.015-0.008-0.017300.1263-0.04710.1140.22450.07310.193519.142485.633774.7804
50.6094-0.2504-0.04210.20230.17390.2133-0.2603-0.2528-0.08260.1508-0.1322-0.06910.0851-0.04570.0002-0.0412-0.3482-0.2528-0.071-0.23690.043113.27676.554594.8623
60.00220.00130.00170.00230.00210.00210.00730.0145-0.0016-0.0205-0.0144-0.0051-0.01350.0105-00.4966-0.1968-0.10780.5194-0.23540.361618.186188.1059113.867
70.01020.0092-0.01010.0042-0.00240.0076-0.03610.010.001-0.023-0.00670.04010.029-0.0062-00.27440.0463-0.16720.53720.02260.14917.068277.7652113.8428
80.02070.0108-0.03870.0146-0.00280.03890.0228-0.01130.1616-0.03740.0373-0.0526-0.0512-0.013900.0627-0.2247-0.18490.1929-0.16850.292615.005783.8332101.1445
9-0.00080.0004-0.00080.00060.0002-0.0003-0.0037-0.0029-0.0080.0022-0.01-0.0025-0.0055-0.006-00.7679-0.1452-0.05570.62970.03410.648110.1735111.943427.3327
100.0039-0.00020.00690.01200.0043-0.00920.03820.0156-0.00010.0240.0223-0.0041-0.002400.3917-0.0965-0.16880.3872-0.05230.2058-2.338112.9150.7283
110.00080.0001-0.00140.00230.00060.0015-0.00890.02580.00860.0284-0.02060.02350.0154-0.030500.2072-0.0591-0.01390.2277-0.00020.2264-12.8485108.884172.9988
120.0007-0.0007-0.00130.00330.00010.0020.0062-0.00540.0032-0.0002-0.00220.00570.0004-0.0022-00.58470.01710.01010.6435-0.01550.5311-16.4284100.962683.6559
130.00040.00090.00140.00070.00110.0015-0.01810.00610.01320.0078-0.0044-0.01020.0072-0.0003-00.3127-0.03850.04990.23020.03270.1471-10.487396.86983.5123
140.00280.0010.00130.0056-0.00150.00380.01130.0057-0.0188-0.00030.01660.00930.01220.0125-00.6268-0.0497-0.07640.5416-0.03010.5947-10.9643104.679385.614
150.00050.0004-0.0007-0.0002-0.00190.0016-0.0181-0.0004-0.00550.01460.029-0.00030.0185-0.002600.2489-0.0554-0.0210.169-0.05150.1349-2.4375109.155874.5964
160.0088-0.00110.00010.00080.00020.0004-0.00310.00140.0001-0.0031-0.00880.00130.00080.0035-00.4243-0.0544-0.03950.45490.02660.47020.5687116.513871.6892
172.00525.82950.12776.1983.02882.97310.256-1.0013-0.94870.43650.3643-0.46750.3854-0.2127-0.63080.0080.0836-0.23890.20340.07560.170237.96453.58689.639
180.00250.0003-0.00040.00040.00050.0001-0.01210.0157-0.0132-0.00750.00290.0004-0.0038-0.010-0.0563-0.0821-0.0750.10920.02150.048630.508664.46194.7302
190.03420.03790.01020.0139-0.0151-0.00530.0750.1235-0.029-0.0977-0.0996-0.07950.05710.17400.1538-0.16320.1982-0.03370.0380.360313.373273.94866.9925
200.0067-0-0.00360.02150.01140.0021-0.0371-0.00420.0062-0.09870.05530.0976-0.05550.025500.3283-0.06940.03190.20960.06880.3149-7.483266.491952.7181
21-0.00030.0060.00270.0053-0.00040.0083-0.02690.0196-0.0357-0.01-0.05770.02040.0287-0.025100.3185-0.0841-0.00160.283-0.00070.2026-3.561589.980857.4799
220.0047-0.0102-0.00250.00750.01820.0164-0.04420.0322-0.0319-0.00940.09780.02930.03470.050400.44710.02910.04010.23770.01190.2091-2.466965.549951.4016
23-0-0.00590.0050.0059-0.00470.01090.04590.0137-0.0449-0.0050.0315-0.0516-0.0081-0.036900.3636-0.05520.03990.179-0.00410.1965-8.417561.558781.6971
240.06260.01660.102-0.0299-0.05830.0144-0.29790.021-0.2016-0.05890.11260.348-0.18320.0793-00.1603-0.134-0.09770.2077-0.05980.1435-3.7597104.569658.2822
250.00120.0009-0.0009-0.00010.00020.0037-0.010.0078-0.00560.01740.00680.00250.0295-0.005800.2257-0.14020.03210.17320.170.083-18.234981.790681.628
260.0044-0.00270.00210.0007-0.00050.0030.0204-0.0321-0.03170.00420.02590.02780.01920.023500.22010.0256-0.01350.183-0.10510.2066-3.581192.506278.4776
270.00190.00050.00210.00410.00720.00410.0087-0.0032-0.00750.00860.0144-0.0262-0.0060.0355-00.3671-0.1997-0.14510.2916-0.09110.16079.2816107.322267.0562
280.0019-0.00030.00230.002100.0051-0.02670.0258-0.0514-0.0081-0.00030.0082-0.0052-0.0006-00.457-0.2605-0.15120.39950.02170.168716.407122.562447.3174
290.0084-0.00530.00810.00670.00680.0152-0.02350.0206-0.00440.01480.01190.00540.01710.0388-00.2618-0.3126-0.03920.379-0.03740.236717.4215109.598852.5319
300.00160.00180.00390.00180.00260.0079-0.04130.00580.00350.0109-0.0606-0.0095-0.00560.0383-00.2764-0.2256-0.01190.2116-0.11910.1086-2.003294.118266.3457
310.0019-0.00080.00070.0033-0.00130.00350.00490.0258-0.0066-0.0026-0.01840.01040.00270.0251-00.0943-0.2023-0.06540.34550.00630.2657-18.530584.790871.4654
320.00110.0006-0.0012-0.0003-0.00060.0009-0.0160.01090.0129-0.0091-0.00760.0144-0.0094-0.0079-00.1057-0.08480.15560.20360.13080.2508-29.808383.707675.3688
330.0177-0.0049-0.01860.02290.00490.0167-0.0097-0.01410.0541-0.0387-0.00060.0558-0.00460.088800.2-0.0679-0.04210.2505-0.31720.2858-9.635581.1591109.3639
340.0009-0.000900.0014-0.00070.00040.00140.00780.00740.0053-0.0133-0.0101-0.00850.012600.43550.07570.00710.2188-0.030.3685-11.718287.267108.1109
35-0.02090.0097-0.01110.00070.0065-0.0086-0.0668-0.26190.0348-0.00610.0677-0.0557-0.0907-0.0886-00.1554-0.09-0.228-0.0401-0.713-0.6585-15.853470.4982119.2503
360.0008-0.00130.00070.00160.00030.00090.004-0.0034-0.00950.00490.0034-0.0098-0.0109-0.0058-00.619-0.04740.0060.6273-0.09490.4872-15.421972.6527135.8079
370.1497-0.05920.10550.25570.04470.0594-0.0489-0.0755-0.1128-0.1161-0.1113-0.09110.03210.13270.0015-0.0053-0.1247-0.49790.2938-0.457-0.6901-1.127357.7992123.5173
380.00330.0006-0.00280.00160.0070.0075-0.0180.0126-0.02350.00020.02470.0450.0087-0.014600.35630.0636-0.36790.497-0.05870.17578.141745.4654123.1179
390.00080.00010.00140.00020.00060.00060.00040.0065-0.0054-0.0004-0.0072-0.00120.0003-0.002200.66760.0678-0.11610.7221-0.09020.72521.793744.7968124.4645
400.001-0.0180.00110.04020.03330.04460.0044-0.1083-0.11220.0923-0.0342-0.03830.0196-0.026200.3370.1935-0.32080.30370.025-0.06680.82451.7904127.7334
41-0.0018-0.0009-0.00220.0008-0.00060.00020.00030.0280.0064-0.0099-0.00210.00870.01410.0175-00.87690.12430.21250.93720.02790.7948-64.796289.0619152.9109
420.00010.0005-0.001-0.0001-0.00010.0048-0.00350.00260.0069-0.00320.00090.0052-0.00740.0012-00.8096-0.05530.08440.7959-0.01690.7528-62.263577.0508147.9954
430.00070.00010.0005-0.0001-0.00020.00110.0110.0110.0044-0.00630.0001-0.00070.00740.000200.76520.01650.11410.8614-0.01750.6993-57.633767.4897144.4714
4400.0003-0.0010.0007-0.00080.00080.00170.0025-0.0106-0.0114-0.0104-0.00090.0159-0.0058-00.6358-0.07260.09250.63150.0480.4788-52.136557.0538140.2708
450.0008-0.00140.00210.0037-0.00210.00270.0231-0.0117-0.0077-0.02360.0082-0.00050.026-0.009-00.5664-0.01230.11840.61520.02070.4911-44.723746.3196129.0625
460.001-0.0012-0.00070.0003-0.00020.0031-0.0068-0.00880.00620.0014-0.00860.00030.0068-0.0007-00.42590.0470.00910.46320.05870.3549-33.680447.1838122.9793
470.0023-0.00150.00090.0026-0.0010.00040.00340.01-0.00330.01530.01110.0049-0.00530.002501.03210.08160.04941.03650.05750.9813-32.964840.9956124.6313
48-0.00050.0003-0.00050.00110.00170.00110.01570.0003-0.009-0.01980.00620.01540.00610.0018-00.7438-0.06880.05480.7485-0.00430.5085-39.291250.3952138.5048
49-0.00040.00560.005-0.0006-0.00040.0038-0.05420.0232-0.0214-0.0105-0.06750.02950.0238-0.0235-00.3767-0.0332-0.1470.508-0.07870.257516.224280.6388121.8339
50-0.02120.0033-0.02510.00180.02240.0276-0.0593-0.08090.1684-0.0386-0.01140.0508-0.09620.046300.3225-0.03040.04130.3387-0.4543-0.2932-24.156982.4973123.6972
510.0218-0.00020.00380.0173-0.0130.0132-0.0563-0.0346-0.0266-0.00770.07190.0047-0.0387-0.081700.18190.01340.10150.529-0.08190.4764-38.050786.605105.4105
520.0047-0.0027-0.0007-0.00020.00230.00630.00570.01370.03170.05070.0474-0.0041-0.0020.0013-00.2822-0.0779-0.00660.587-0.09320.4279-43.420972.8859123.7732
530.01050.00050.01440.01540.00330.011-0.08130.02530.0366-0.0505-0.0682-0.0277-0.045-0.038100.2410.14670.11230.4734-0.07880.321-36.319490.1091108.3567
540.0202-0.01810.02010.01840.00020.0136-0.0281-0.0359-0.12670.05850.1630.09550.07040.0693-00.1995-0.0267-0.03330.2683-0.13560.1529-26.110958.8661100.7174
550.03130.01220.0060.0178-0.01640.0110.0861-0.05640.02370.0834-0.00670.057-0.0453-0.003300.3105-0.09020.45860.5649-0.19150.1254-52.137271.3175145.7336
56-00.0004-0.0022-0.0001-0.00050.00060.0146-0.00220.00470.01210.01160.01-0.0092-0.002700.82720.00290.05830.798-0.06210.737-56.63161.9564153.686
570.0008-0.0010.0011-0.0001-0.00160.0020.0009-0.00820.006-0.0061-0.0060.0037-0.00110.010800.2696-0.0720.08360.3794-0.01070.2949-32.90651.8762104.4
580.00450.00050.00240.00490.00140.00210.0036-0.026-0.00720.03460.02840.002-0.01520.0097-00.398-0.0718-0.01190.430.01790.2787-30.486755.7455122.5684
590.00060.00050.00110.0006-0.00020.0010.001-0.01780.00690.0001-0.0140.0023-0.00170.001700.5907-0.0365-0.00870.6734-0.09190.4042-34.178860.4573141.1531
600.00050.00030.0003-0.00010.00060.0010.00550.0012-0.00230.005-0.0062-0.0067-0.001-0.001800.64130.03170.07440.7227-0.05310.582-39.349764.6744152.4697
610.00080.00040.0004-0.00130.00030.0004-0.00410.0124-0.0079-0.00710.0031-0.001-0.0073-0.005-00.8017-0.03010.00830.968-0.0980.7469-49.607469.8744161.8461
620.0001-0.00020.0004-0.00080.00050.0015-0.0077-0.0044-0.0026-0.0003-0.0177-0.0040.0025-0.0029-01.25540.11080.06951.2861-0.04531.0319-50.78378.0439166.0973
630.01090.011-0.00240.0103-0.00630.00250.0256-0.05020.0075-0.03980.0330.04830.02880.004100.3809-0.0119-0.07640.502-0.17280.1003-39.311769.5686139.9099
640.006-0.0032-0.0060.00610.00240.0027-0.0462-0.0023-0.0021-0.0429-0.0507-0.03150.0004-0.006400.3673-0.00040.05640.2764-0.03490.2753-41.74755.9177106.9371
650.00080.0020.00070.00020.00190.0024-0.02350.00920.0050.01980.01730.00680.0077-0.0087-00.1526-0.0095-0.02240.19550.01840.1837-23.647341.9309105.1118
660.0040.00330.00220.0053-0.00110.0081-0.0153-0.01330.00560.0173-0.0067-0.00640.0298-0.0169-00.23980.03960.06090.2160.05910.3262-5.363734.0434110.4327
670.0934-0.00430.05790.04140.04490.09310.2372-0.8297-0.07840.6339-0.1162-0.15950.48220.1561-0-0.82491.1182-0.1277-1.02010.2325-0.0037-0.876130.04995.4658
680.15090.0420.0050.0978-0.11070.1553-0.0137-0.06980.077-0.08570.1785-0.0738-0.16330.1591-0.0067-0.24320.40990.0249-0.04120.06570.443616.088141.548994.7492
690.00470.00080.00170.0113-0.00720.0047-0.01040.0046-0.0179-0.02520.0323-0.02110.012-0.010700.3179-0.0657-0.00970.23070.11360.191827.07851.469592.1637
700.0143-0.0050.0120.00450.00390.0075-0.0611-0.01940.02460.0126-0.0322-0.00350.06010.0662-00.2575-0.104-0.29520.36020.19350.549938.116149.428699.3743
710.0036-0.0014-0.00170.00240.00540.00110.0036-0.0161-0.0319-0.0409-0.0543-0.01950.01730.029900.17330.0438-0.08020.3683-0.09250.305123.441429.904390.9524
720.0014-0.00040.00110.0028-0.00840.0085-0.0316-0.01920.027-0.0458-0.06540.07750.0227-0.0165-00.3126-0.0159-0.01590.22790.16920.1857-1.153243.723679.0121
730.0005-0.001100.00040.0002-0.00010.0037-0.0022-0.00030.01510.0053-0.00220.0033-0.0003-00.45030.0202-0.02840.4528-0.08070.4741-17.8303-29.547877.7793
740.00010.0002-00.001-0.0009-0.0002-0.00790.003-0.0064-0.0036-0.0008-0.00860.0049-0.003-00.4813-0.02440.05960.35640.0210.5466-15.8972-22.42571.7676
750.00320.00140.00440.00390.00460.00480.00390.00050.0187-0.02340.0264-0.0141-0.05930.0295-00.2961-0.008-0.07950.2204-0.09540.3006-10.0847-5.42763.9893
760.0016-0.00090.00130.00170.00150.0013-0.0210.02040.00360.001-0.02880.00530.03240.0362-00.18870.0318-0.04850.1333-0.08980.2427-3.486814.232957.4804
770.0011-0.002-0.00070.0001-0.00320.00270.00780.01830.0032-0.04190.020.01520.0236-0.013200.34030.04670.09710.3301-0.05880.3167-2.212728.479159.6728
780.00160.0002-0.00020.0004-0.00050.0002-0.0189-0.0094-0.0008-0.0144-0.00020.016-0.01620.001100.3302-0.05140.04990.3843-0.11180.4730.134136.80865.3773
790.00130.00030.00040.00190.00060.0006-0.00210.00220.00240.0046-0.0098-0.00360.00310.0054-00.3006-0.00560.08860.383-0.06270.29086.978422.578159.1984
800.00160.00010.00010.0007-0.00120.00210.01790.01140.01370.00580.0241-0.00410.02520.015600.25730.09460.04420.1486-0.02550.14683.487413.415364.4679
810.00480.0054-0.00220.0038-0.00240.0048-0.0345-0.04550.03150.0057-0.0271-0.0022-0.00260.087500.35680.03660.02020.1633-0.06640.2057-8.287537.040373.5383
820.0044-0.0002-0.00120.00150.0030.00780.0225-0.01340.0273-0.00580.0084-0.0105-0.0037-0.0356-00.17280.07670.00520.23360.03550.27224.842310.941973.1927
830.0027-0.00150.00120.00170.00090.0009-0.02220.0006-0.01640.0064-0.00710.00530.01370.018600.31820.155-0.11980.17450.03980.48056.7279-10.535170.4702
840.00180.00150.00180.00210.00130.0018-0.0325-0.01170.0012-0.0023-0.0143-0.00220.00050.002400.3796-0.03070.00690.19090.08690.32924.5258-22.481165.2321
850.0009-0.0005-0.00040.0026-0.00150.00060.0169-0.00380.0077-0.00340.0182-0.0050.007-0.006700.1874-0.083-0.04860.1953-0.02020.33914.4815-25.175373.4446
860.0027-0.0020.0010.0009-0.00010.0013-0.039-0.0083-0.01310.0154-0.0154-0.0014-0.00710.011-00.32210.1001-0.08230.2666-0.05670.45434.0275-14.218579.3622
870.0021-0.00040.00180.00220.00210.00160.0168-0.0042-0.00610.0069-0.0096-0.0164-0.01060.0017-00.30560.1348-0.15050.2076-0.0450.1551-0.39960.965278.4611
880.01110.01280.00540.0094-0.0006-0.0008-0.05940.04950.0136-0.03580.08420.05-0.0644-0.0306-00.13030.20610.16990.0842-0.00540.4284-15.192129.250671.0998
89-0.0010.00040.002-0.00090.0012-0.00130.0207-0.01130.0049-0.0210.00950.00490.0052-0.001400.48910.0768-0.06180.56540.05760.527817.023339.1681132.8235
900.0010.00010.00190.0001-00.00040.0033-0.01240.00190.00640.01280.00560.0032-0.0079-00.3944-0.0124-0.1810.32280.06520.295414.792637.172117.873
910.0059-0.0056-0.0068-0.0035-0.0090.02130.0548-0.1257-0.0870.0752-0.0451-0.0460.10610.0768-00.36770.05420.0734-0.14130.24670.1481-5.936621.1846100.6149
920.02130.00280.01220.0109-0.00010.01790.0090.0304-0.0438-0.0188-0.0540.03410.0007-0.135500.29260.00080.03690.22160.01860.3536-31.344223.33894.0077
930.00450.0026-0.00240.0072-0.00320.0029-0.01920.0001-0.00370.0672-0.0372-0.00250.0014-0.0273-00.217-0.007-0.01440.2496-0.05010.4127-15.966514.051877.9698
940.00850.0022-0.00390.0112-0.01930.01730.0537-0.0205-0.0293-0.00320.00740.0301-0.03430.01400.36430.0064-0.07390.1794-0.03290.3411-29.645618.811197.1281
950.0023-0.00560.01110.00240.0080.0127-0.0267-0.11480.0204-0.06050.08490.06440.0297-0.0301-00.21240.1220.00520.2769-0.00410.2087-16.042745.801194.6986
960.08040.00160.04070.00510.01510.03260.0240.119-0.21520.03860.0536-0.0238-0.0417-0.0715-00.08830.1825-0.0698-0.25260.09320.4036-8.87626.932267.5219
Refinement TLS group
IDRefine-IDRefine TLS-IDSelection details
1X-RAY DIFFRACTION1(chain A and resid 154:187)
2X-RAY DIFFRACTION2(chain A and resid 188:232)
3X-RAY DIFFRACTION3(chain A and resid 233:277)
4X-RAY DIFFRACTION4(chain A and resid 278:284)
5X-RAY DIFFRACTION5(chain A and resid 285:356)
6X-RAY DIFFRACTION6(chain A and resid 357:372)
7X-RAY DIFFRACTION7(chain A and resid 373:397)
8X-RAY DIFFRACTION8(chain A and resid 398:445)
9X-RAY DIFFRACTION9(chain B and resid 1:11)
10X-RAY DIFFRACTION10(chain B and resid 12:34)
11X-RAY DIFFRACTION11(chain B and resid 35:45)
12X-RAY DIFFRACTION12(chain B and resid 46:68)
13X-RAY DIFFRACTION13(chain B and resid 69:75)
14X-RAY DIFFRACTION14(chain B and resid 76:89)
15X-RAY DIFFRACTION15(chain B and resid 90:101)
16X-RAY DIFFRACTION16(chain B and resid 102:106)
17X-RAY DIFFRACTION17(chain C and resid 63:72)
18X-RAY DIFFRACTION18(chain C and resid 73:77)
19X-RAY DIFFRACTION19(chain C and resid 118:179)
20X-RAY DIFFRACTION20(chain C and resid 180:227)
21X-RAY DIFFRACTION21(chain C and resid 231:259)
22X-RAY DIFFRACTION22(chain C and resid 260:293)
23X-RAY DIFFRACTION23(chain C and resid 294:321)
24X-RAY DIFFRACTION24(chain C and resid 322:395)
25X-RAY DIFFRACTION25(chain D and resid 1:11)
26X-RAY DIFFRACTION26(chain D and resid 12:23)
27X-RAY DIFFRACTION27(chain D and resid 24:41)
28X-RAY DIFFRACTION28(chain D and resid 42:61)
29X-RAY DIFFRACTION29(chain D and resid 62:84)
30X-RAY DIFFRACTION30(chain D and resid 85:99)
31X-RAY DIFFRACTION31(chain D and resid 100:111)
32X-RAY DIFFRACTION32(chain D and resid 112:120)
33X-RAY DIFFRACTION33(chain E and resid 153:186)
34X-RAY DIFFRACTION34(chain E and resid 187:193)
35X-RAY DIFFRACTION35(chain E and resid 194:277)
36X-RAY DIFFRACTION36(chain E and resid 278:282)
37X-RAY DIFFRACTION37(chain E and resid 283:366)
38X-RAY DIFFRACTION38(chain E and resid 367:391)
39X-RAY DIFFRACTION39(chain E and resid 392:396)
40X-RAY DIFFRACTION40(chain E and resid 397:445)
41X-RAY DIFFRACTION41(chain F and resid 1:11)
42X-RAY DIFFRACTION42(chain F and resid 12:17)
43X-RAY DIFFRACTION43(chain F and resid 18:25)
44X-RAY DIFFRACTION44(chain F and resid 26:34)
45X-RAY DIFFRACTION45(chain F and resid 35:66)
46X-RAY DIFFRACTION46(chain F and resid 67:74)
47X-RAY DIFFRACTION47(chain F and resid 75:89)
48X-RAY DIFFRACTION48(chain F and resid 90:102)
49X-RAY DIFFRACTION49(chain G and resid 60:125)
50X-RAY DIFFRACTION50(chain G and resid 126:179)
51X-RAY DIFFRACTION51(chain G and resid 180:224)
52X-RAY DIFFRACTION52(chain G and resid 225:259)
53X-RAY DIFFRACTION53(chain G and resid 260:292)
54X-RAY DIFFRACTION54(chain G and resid 293:340)
55X-RAY DIFFRACTION55(chain G and resid 341:390)
56X-RAY DIFFRACTION56(chain G and resid 391:399)
57X-RAY DIFFRACTION57(chain H and resid 1:8)
58X-RAY DIFFRACTION58(chain H and resid 9:24)
59X-RAY DIFFRACTION59(chain H and resid 25:34)
60X-RAY DIFFRACTION60(chain H and resid 35:41)
61X-RAY DIFFRACTION61(chain H and resid 42:54)
62X-RAY DIFFRACTION62(chain H and resid 55:69)
63X-RAY DIFFRACTION63(chain H and resid 70:98)
64X-RAY DIFFRACTION64(chain H and resid 99:120)
65X-RAY DIFFRACTION65(chain I and resid 154:162)
66X-RAY DIFFRACTION66(chain I and resid 163:192)
67X-RAY DIFFRACTION67(chain I and resid 193:285)
68X-RAY DIFFRACTION68(chain I and resid 286:366)
69X-RAY DIFFRACTION69(chain I and resid 367:393)
70X-RAY DIFFRACTION70(chain I and resid 394:416)
71X-RAY DIFFRACTION71(chain I and resid 417:430)
72X-RAY DIFFRACTION72(chain I and resid 431:445)
73X-RAY DIFFRACTION73(chain J and resid 1:5)
74X-RAY DIFFRACTION74(chain J and resid 6:12)
75X-RAY DIFFRACTION75(chain J and resid 13:30)
76X-RAY DIFFRACTION76(chain J and resid 31:41)
77X-RAY DIFFRACTION77(chain J and resid 42:70)
78X-RAY DIFFRACTION78(chain J and resid 71:86)
79X-RAY DIFFRACTION79(chain J and resid 87:93)
80X-RAY DIFFRACTION80(chain J and resid 94:102)
81X-RAY DIFFRACTION81(chain K and resid 1:20)
82X-RAY DIFFRACTION82(chain K and resid 21:39)
83X-RAY DIFFRACTION83(chain K and resid 40:49)
84X-RAY DIFFRACTION84(chain K and resid 50:57)
85X-RAY DIFFRACTION85(chain K and resid 58:64)
86X-RAY DIFFRACTION86(chain K and resid 65:74)
87X-RAY DIFFRACTION87(chain K and resid 75:85)
88X-RAY DIFFRACTION88(chain K and resid 86:120)
89X-RAY DIFFRACTION89(chain N and resid 63:69)
90X-RAY DIFFRACTION90(chain N and resid 70:76)
91X-RAY DIFFRACTION91(chain N and resid 118:179)
92X-RAY DIFFRACTION92(chain N and resid 180:226)
93X-RAY DIFFRACTION93(chain N and resid 227:257)
94X-RAY DIFFRACTION94(chain N and resid 258:289)
95X-RAY DIFFRACTION95(chain N and resid 290:321)
96X-RAY DIFFRACTION96(chain N and resid 322:395)

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