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- PDB-5j9u: Crystal structure of the NuA4 core complex -

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Basic information

Entry
Database: PDB / ID: 5j9u
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
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 / SUMOylation of transcription cofactors ...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 / SUMOylation of transcription cofactors / positive regulation of triglyceride biosynthetic process / DNA-templated transcription elongation / histone H4 acetyltransferase activity / rDNA heterochromatin formation / protein N-acetyltransferase activity / Recruitment and ATM-mediated phosphorylation of repair and signaling proteins at DNA double strand breaks / protein-lysine-acetyltransferase activity / NuA4 histone acetyltransferase complex / histone acetyltransferase activity / Estrogen-dependent gene expression / positive regulation of macroautophagy / histone H4K16 acetyltransferase activity / histone H3K56 acetyltransferase activity / histone H3K23 acetyltransferase activity / histone H2AK5 acetyltransferase activity / histone H2AK9 acetyltransferase activity / histone H2BK5 acetyltransferase activity / histone H2BK12 acetyltransferase activity / histone H3K4 acetyltransferase activity / histone H3K27 acetyltransferase activity / histone H3K36 acetyltransferase activity / histone H3K122 acetyltransferase activity / histone H3K18 acetyltransferase activity / histone H3K9 acetyltransferase activity / histone H3K14 acetyltransferase activity / histone H4K5 acetyltransferase activity / histone H4K8 acetyltransferase activity / histone H4K12 acetyltransferase activity / histone acetyltransferase / Transferases; Acyltransferases; Transferring groups other than aminoacyl groups / methylated histone binding / meiotic cell cycle / positive regulation of transcription elongation by RNA polymerase II / transcription coregulator activity / nucleosome / regulation of cell cycle / chromatin remodeling / DNA repair / DNA-templated transcription / chromatin binding / regulation of DNA-templated transcription / regulation of transcription by RNA polymerase II / chromatin / nucleus / metal ion binding / 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 / Chromo-like domain superfamily / Enhancer of polycomb-like, N-terminal / Enhancer of polycomb-like / 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 / Winged helix-like DNA-binding domain superfamily/Winged helix DNA-binding domain / Zinc finger, FYVE/PHD-type / 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
Similarity search - Component
Biological speciesSaccharomyces cerevisiae (brewer's yeast)
MethodX-RAY DIFFRACTION / SYNCHROTRON / MOLECULAR REPLACEMENT / Resolution: 2.95 Å
AuthorsChen, Z.C. / Xu, P.
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 23, 2024Group: Data collection / Database references ...Data collection / Database references / Derived calculations / Structure summary
Category: chem_comp_atom / chem_comp_bond ...chem_comp_atom / chem_comp_bond / citation / database_2 / pdbx_entry_details / pdbx_modification_feature / pdbx_struct_oper_list
Item: _citation.journal_id_CSD / _database_2.pdbx_DOI ..._citation.journal_id_CSD / _database_2.pdbx_DOI / _database_2.pdbx_database_accession / _pdbx_struct_oper_list.symmetry_operation

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Structure visualization

Structure viewerMolecule:
MolmilJmol/JSmol

Downloads & links

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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


Theoretical massNumber of molelcules
Total (without water)315,41312
Polymers315,41312
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


Theoretical massNumber of molelcules
Total (without water)105,1384
Polymers105,1384
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area18610 Å2
ΔGint-104 kcal/mol
Surface area41700 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


Theoretical massNumber of molelcules
Total (without water)105,1384
Polymers105,1384
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area18200 Å2
ΔGint-105 kcal/mol
Surface area41540 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


Theoretical massNumber of molelcules
Total (without water)105,1384
Polymers105,1384
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area18150 Å2
ΔGint-105 kcal/mol
Surface area41470 Å2
MethodPISA
Unit cell
Length a, b, c (Å)217.909, 189.889, 173.558
Angle α, β, γ (deg.)90.00, 113.61, 90.00
Int Tables number5
Space group name H-MC121

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Components

#1: Protein Histone acetyltransferase ESA1


Mass: 36557.867 Da / Num. of mol.: 3 / Fragment: UNP residues 141-445)
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
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: 5.22 Å3/Da / Density % sol: 76.42 %
Crystal growTemperature: 298 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: Apr 20, 2014
RadiationProtocol: MAD / Monochromatic (M) / Laue (L): M / Scattering type: x-ray
Radiation wavelengthWavelength: 0.987 Å / Relative weight: 1
ReflectionResolution: 2.95→30 Å / Num. obs: 106917 / % possible obs: 78.4 % / Redundancy: 2.8 % / Net I/σ(I): 10.3

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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: 2.95→29.686 Å / SU ML: 0.37 / Cross valid method: FREE R-VALUE / σ(F): 1.41 / Phase error: 27.94
RfactorNum. reflection% reflection
Rfree0.2462 1993 1.86 %
Rwork0.2157 --
obs0.2162 106917 78.84 %
Solvent computationShrinkage radii: 0.9 Å / VDW probe radii: 1.11 Å
Refinement stepCycle: LAST / Resolution: 2.95→29.686 Å
ProteinNucleic acidLigandSolventTotal
Num. atoms19676 0 0 0 19676
Refine LS restraints
Refine-IDTypeDev idealNumber
X-RAY DIFFRACTIONf_bond_d0.00320089
X-RAY DIFFRACTIONf_angle_d0.69427054
X-RAY DIFFRACTIONf_dihedral_angle_d14.64312290
X-RAY DIFFRACTIONf_chiral_restr0.0432902
X-RAY DIFFRACTIONf_plane_restr0.0043485
LS refinement shell
Resolution (Å)Rfactor RfreeNum. reflection RfreeRfactor RworkNum. reflection RworkRefine-ID% reflection obs (%)
2.95-3.02370.3552450.29672362X-RAY DIFFRACTION25
3.0237-3.10540.3007720.2913831X-RAY DIFFRACTION40
3.1054-3.19670.3447940.29754946X-RAY DIFFRACTION52
3.1967-3.29970.29261160.29256093X-RAY DIFFRACTION64
3.2997-3.41750.34751400.28477389X-RAY DIFFRACTION78
3.4175-3.55410.31811640.26978633X-RAY DIFFRACTION91
3.5541-3.71560.31761710.24559005X-RAY DIFFRACTION94
3.7156-3.91110.25011720.22759027X-RAY DIFFRACTION95
3.9111-4.15550.23681710.21319001X-RAY DIFFRACTION95
4.1555-4.47540.21521700.18958958X-RAY DIFFRACTION94
4.4754-4.92390.22891700.17938927X-RAY DIFFRACTION94
4.9239-5.63220.22271700.19339003X-RAY DIFFRACTION95
5.6322-7.080.22391710.22528969X-RAY DIFFRACTION94
7.08-29.68720.19361670.17058780X-RAY DIFFRACTION91
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)
13.00692.7510.24675.36990.36192.17750.01370.4223-0.2261-0.37960.1412-0.40060.33050.0845-0.12080.38060.06030.07920.31960.01070.481910.620959.465976.184
21.3929-0.4829-0.22982.00131.5224.353-0.02640.3370.0165-0.41720.0823-0.42480.14930.1527-0.07410.2119-0.06230.09730.23820.0420.362511.22374.202571.1775
32.06270.3242-0.39742.37420.15781.4027-0.0667-0.20430.03120.0792-0.05050.10430.1275-0.03130.08180.1442-0.04440.01480.22590.04620.28955.181573.700884.7501
44.62123.69448.06288.37996.08162.00090.09870.10250.1474-0.4921-0.1202-1.0879-0.44141.77150.03120.2902-0.10530.26660.48010.17260.785718.815485.686175.2348
53.0757-0.2232-0.68331.89480.41712.9358-0.1565-0.5492-0.27310.2342-0.0741-0.2353-0.020.11970.21940.2455-0.0442-0.05470.26020.01150.395413.002376.948495.5073
65.5765-1.2371-4.50033.1540.09735.47480.233-0.75690.95630.4984-0.08390.1671-0.7180.2205-0.14940.74-0.1883-0.21770.7758-0.16590.67717.887188.779114.4374
74.7996-1.8330.19133.7921-2.45735.004-0.1044-0.48330.01070.4736-0.0071-0.46540.04090.43170.11570.6185-0.0756-0.21460.7397-0.01910.540916.774178.6229114.4927
81.9150.04360.20980.6254-0.54251.84110.1421-0.13660.15160.3529-0.0806-0.3042-0.12750.5298-0.12620.4848-0.0452-0.11180.4378-0.13490.702914.721584.3905101.7375
91.66841.5415-0.06632.3529-0.81580.6162-0.02610.7108-0.2932-0.53280.0037-0.2630.17960.02310.02351.3893-0.3402-0.1251.1442-0.06780.87299.6205111.893827.974
103.45161.5561-5.40611.1184-3.45172.0060.03240.50410.2847-0.19120.010.1737-0.3273-0.2011-0.11440.6106-0.1502-0.23920.60050.01630.5328-2.6522112.794951.121
118.42542.1828-2.3347.3554-0.63670.64490.0495-0.070.46680.2399-0.05090.3644-0.4415-0.182-0.02440.50410.02940.08590.2450.0480.5907-13.2629108.761973.419
128.76011.5167-6.2851.59321.44289.3217-0.0331-0.17760.51630.207-0.02520.7202-0.5991-0.70650.09010.8161-0.0266-0.10870.9321-0.09120.7694-16.7295100.702984.0851
132.00265.5006-0.56762.0013-3.23112.00180.3879-1.2183-0.23271.5299-0.38550.37380.0414-0.04060.00130.688-0.1346-0.07920.5722-0.06090.4396-10.677196.649583.8105
140.00060.00230.01120.00580.03120.14920.2977-0.4849-0.12610.8514-0.4676-0.2171-0.24740.2050.14760.8341-0.085-0.05370.8606-0.27790.9974-11.3245104.461886.1924
158.05592.3954-2.39158.1896-4.48475.28850.008-0.53250.48410.4105-0.0174-0.018-0.59130.2379-0.03410.47570.00060.01290.1823-0.1050.565-2.8597108.967175.0078
168.3354.5893-5.97822.7721-2.91044.8790.3446-0.10472.1158-0.0001-0.1571-0.6927-1.86120.8363-0.20521.001-0.1081-0.01790.649-0.05091.10430.0368116.461272.0426
172.39371.4518-1.26832.94920.26551.18190.34530.1771-0.20340.3422-0.3463-0.85980.36890.91560.09680.60420.0809-0.26420.77760.08531.049440.586352.848192.4373
182.00182.00091.99982.00311.99952.00020.4084-0.3628-0.76690.6674-0.03990.29140.71890.0872-0.35790.4102-0.0589-0.08510.64520.19660.869330.223364.606395.0482
191.0060.09370.22961.02640.45310.57690.05320.2017-0.005-0.3711-0.049-0.5075-0.02460.28080.01540.3793-0.03790.13620.32960.04410.57513.105773.922167.6454
203.21522.0326-0.69444.4599-2.00332.3944-0.10790.77990.2273-0.53850.20780.365-0.0240.0295-0.06110.5884-0.0379-0.04930.5545-0.0070.3891-7.679166.511453.3889
216.63670.9653.73254.8945-1.58958.24180.26581.1936-1.0668-1.097-0.1777-0.22651.02570.4499-0.05370.59180.01550.02350.5105-0.14170.745-3.830789.958857.9971
221.3934-0.43210.06375.3367-2.34573.9979-0.07240.41610.1173-0.78070.29230.08520.00310.1089-0.14720.6899-0.05570.02220.4674-0.03290.3516-2.609365.581352.0821
234.1604-1.9854-0.57892.70920.19351.25620.05750.003-0.0820.0974-0.05950.6733-0.0287-0.4085-0.00140.3726-0.04650.0450.39960.00680.7129-8.698861.754282.4344
242.88231.4964-1.96872.0908-1.55463.0124-0.08250.32740.0147-0.25830.07420.1419-0.4030.0249-0.0150.4163-0.0781-0.11120.3754-0.00510.2862-3.991104.479458.7641
252.00149.5911-0.77282.0018-0.65418.15990.1582-0.189-1.17950.3168-0.0658-0.19320.8211-0.15-0.09040.2594-0.0309-0.02740.36150.05280.4499-18.544381.732881.9085
267.00331.7627-1.48731.2938-2.16834.0880.0251-0.2922-0.32040.2039-0.0388-0.09540.0025-0.01840.0280.23830.00610.01010.28330.01630.4924-3.953892.440378.7595
278.36256.852-4.91266.3784-3.69133.02020.1636-0.17280.09790.313-0.1943-0.1342-0.47950.403-0.01360.5269-0.1536-0.13430.36310.01370.54948.9701107.213667.3915
281.0427-0.54320.9955.1604-2.47115.0842-0.01970.1072-0.01050.00670.09350.214-0.0669-0.1767-0.08070.4956-0.1921-0.02870.5470.1050.485516.2406122.488647.7792
293.38274.3662-2.84467.1181-4.50083.0112-0.00260.022-0.1907-0.1359-0.0377-0.3102-0.08380.44360.04630.4611-0.116-0.03510.55860.01480.285717.1268109.532452.9917
301.9997-1.1480.26485.68020.26235.18210.0344-0.0574-0.25350.0162-0.0602-0.2244-0.0240.14070.04910.2713-0.1976-0.09810.2863-0.00490.3342-2.324894.06966.6852
312.00369.902-0.72638.78440.38951.5544-0.1460.4089-0.2757-0.3410.038-0.23450.23580.09980.05130.2524-0.02530.05660.29310.02920.3956-18.895784.733171.8825
322.00324.70370.24127.3849-1.03749.89420.10370.2225-0.241-0.0222-0.11890.39830.1148-0.22250.0610.17840.01860.14860.49350.09880.7216-30.044483.530775.7983
331.6147-1.35650.31252.3421-2.08067.43590.0858-0.23050.4140.219-0.0071-0.0186-0.3443-0.0304-0.0550.394-0.04850.06140.4111-0.1210.4684-10.021581.1316110.0685
341.40281.37871.31352.00041.99982.001-0.33730.24370.8293-0.79780.27040.0152-1.34650.46130.09820.918-0.0761-0.08270.7256-0.02540.555-12.165687.1029108.7162
350.5554-0.11510.06561.3967-0.23830.59790.2194-0.46880.25060.35950.0154-0.3107-0.3102-0.0159-0.00710.4995-0.0365-0.16580.5356-0.5168-1.3721-16.331970.4962119.9133
361.99981.9985-0.0442.0001-6.59362.0015-0.3211-1.50330.3311.020.6402-0.63250.22860.621-0.33190.8628-0.0499-0.09410.8198-0.13910.223-16.134972.6744136.2798
371.7713-0.42430.1541.72930.10221.3222-0.0316-0.5305-0.18530.3512-0.046-0.54190.48910.5839-0.05230.48990.0057-0.26410.8337-0.0519-0.036-1.670557.9014124.423
388.1274-0.62471.62971.32781.87333.41780.0839-0.3386-0.36540.1990.0568-0.08160.33570.1978-0.03030.62730.0675-0.2210.65130.02570.47827.617245.5925124.1949
395.8074-8.9725-2.90682.00038.9236.6998-0.2425-0.49070.08190.54570.738-1.71170.38021.2974-0.47710.92590.1297-0.31911.0536-0.0991.120721.088445.0962125.6618
401.9588-0.0890.36060.63430.66521.1577-0.1533-0.5025-0.04220.33820.1712-0.24050.12930.28440.03530.91120.0314-0.24240.88050.16130.53990.228651.9313128.5941
413.1684-3.4181-1.92083.682.06991.16440.0668-0.04180.4825-0.0576-0.05660.1474-0.6212-0.2697-0.07321.3660.03210.44721.1470.00750.4901-65.394288.8164153.2839
428.9359-6.0212.3152.00147.9142.0007-0.09720.19950.2554-0.0016-0.08560.449-0.3629-0.38830.16030.8424-0.09560.34961.0722-0.01660.6029-62.918177.1767148.2937
435.7744-4.2318-2.32214.38743.48613.40570.04690.0852-0.33430.062-0.02310.50440.4005-0.59920.01930.9101-0.19380.39490.96790.07680.3565-58.3267.4185144.8865
441.85353.35512.55942.00112.71054.2944-0.1752-0.5315-0.14610.5582-0.06640.32160.2247-0.09490.17050.8194-0.17180.18750.96150.25450.3697-52.761157.0415140.4996
451.78230.72430.57065.62570.81017.1704-0.04180.0159-0.41260.04-0.0388-0.06450.59280.21260.15090.79780.0210.18250.75380.24250.4361-45.348546.3246129.3303
462.00082.11440.05592.0011-6.05382.0009-0.2859-0.2419-0.7457-0.0728-0.0661-0.38581.05670.59910.34660.65280.07890.10990.73480.32440.4489-34.162947.2227123.3677
470.0001-0.0008-0.00030.0014-0.00020-0.1668-0.2692-0.33210.0356-0.1219-0.11960.60060.81450.30611.31020.23050.161.23180.34140.8923-33.592940.9947124.9481
485.05051.0589-3.35620.80641.47872.002-0.0906-0.3753-0.38050.32560.2072-0.09720.49590.2932-0.08571.1057-0.17910.00150.93650.1540.2596-39.912750.2631138.8202
493.57752.35780.53172.3251-0.01910.703-0.1298-0.23510.19410.1362-0.1917-0.2049-0.2150.22430.17390.8696-0.0404-0.29691.1017-0.10340.66717.437681.2954123.24
500.24080.25560.03391.0697-0.23280.7666-0.0092-0.4340.31160.33850.1149-0.0416-0.3856-0.0702-0.06630.7388-0.01560.03720.7699-0.17220.3865-24.62782.5062124.2202
511.52050.1973-0.58520.76111.6026.69760.0088-0.15980.29540.09050.02910.2687-0.1892-0.81630.02770.43980.02380.02810.75680.04050.5544-38.293986.6353105.8932
521.4222-1.82831.92592.3426-2.4712.5744-0.0715-0.05490.74050.24830.08230.1379-0.5540.0386-0.01690.7936-0.03990.06390.9683-0.08540.4718-43.997672.8859124.2059
534.4707-0.6118-1.62512.45390.43484.0548-0.0746-0.25930.27360.4272-0.05290.4464-0.2933-0.86380.07490.54730.11550.12580.63080.0070.5693-36.539290.0909108.9012
541.92511.99950.12312.1980.154-0.0071-0.27010.2528-0.0714-0.27390.425-0.0343-0.0106-0.0019-0.12640.3858-0.04110.00040.4065-0.02670.4116-26.379158.9473101.2835
551.3989-2.1595-2.27343.75723.74324.27540.2162-0.4540.3150.546-0.11420.417-0.2859-0.3059-0.01060.9674-0.30870.74010.9790.0211-0.7737-52.890371.3068146.1344
560.26891.37930.29536.77761.490.32580.29730.4485-0.4464-0.5741-0.17390.3888-0.0645-0.1407-0.00661.1475-0.05210.30311.2339-0.06130.526-57.286761.9551154.0028
574.14543.52440.54315.4101-4.32272.0003-0.03330.83980.3017-0.567-0.1258-0.4726-0.2510.8310.12290.5767-0.01940.20350.51350.00410.4262-33.491552.2007104.7471
581.71060.30212.49491.74783.21858.1797-0.03460.0154-0.0004-0.08730.0705-0.1135-0.31110.4239-0.03180.6124-0.0853-0.01750.62030.12210.3226-31.032555.9406123.029
591.30972.05624.30313.3146.48662.0014-0.1334-0.49870.11410.36220.0997-0.2527-0.00570.41920.01860.93010.0090.00580.91050.10220.1886-34.892760.4327141.658
601.03971.83563.14443.31256.16152.0007-0.0555-0.57620.19160.5530.0164-0.5033-0.18590.23820.05460.97950.01070.05110.9774-0.00660.3665-39.963864.7693152.811
617.8920.0308-5.38660.284-0.28643.9241-0.15980.0199-0.35920.08250.0190.21840.3972-0.27430.1381.2686-0.09410.22871.454-0.0340.3156-50.296669.6181162.2167
621.2228-1.807-0.42482.86950.84530.37140.00380.07580.0004-0.0968-0.0316-0.11490.04670.1058-0.01831.48460.11130.20871.5353-0.13540.1809-51.630377.9417166.4788
631.0073-0.6281-0.86621.34650.62290.740.033-0.58370.25510.49460.0742-0.032-0.05960.2692-0.1130.9584-0.25810.11351.0145-0.0644-0.0065-39.985369.6012140.4357
644.78371.79442.35045.5143.22538.9642-0.14150.02360.1386-0.3092-0.10580.2109-0.352-0.07130.18670.35910.05730.04190.36080.00050.2647-42.259756.1468107.358
654.13923.36365.34143.15874.57577.02060.05930.4415-0.2477-0.40060.08690.2594-0.0377-0.0808-0.13580.6979-0.08980.1140.5296-0.08640.7506-23.913342.2414105.8239
662.8497-1.33611.1445.16290.05341.8729-0.1861-0.8226-0.39140.68750.18260.220.2781-0.21080.03720.48550.04460.02850.41010.06710.4386-5.607934.37111.0008
671.28850.2521-0.47721.5055-0.14921.3378-0.0544-0.1681-0.23630.1928-0.0417-0.04290.12150.08620.05320.28920.0814-0.02240.22240.03840.4742-1.171530.408296.0879
682.28480.37410.22841.4531-0.30981.4544-0.22120.00780.11040.098-0.0807-0.6134-0.12550.26070.23660.2710.052-0.02250.27610.04560.586215.879541.729295.2698
695.62970.5443.1025.26351.32715.4177-0.0743-0.10480.1556-0.0798-0.081-0.1789-0.1035-0.03030.07790.2805-0.0285-0.01410.28130.10050.685126.949351.574592.6896
700.94430.86110.10945.2647-0.44611.14280.0077-0.24340.15920.5779-0.1469-0.4135-0.02020.44510.12530.4791-0.0023-0.26310.6830.17151.480537.921949.428899.7448
712.0048-2.43575.71332.1629-3.34935.78130.0065-0.3562-0.32750.21980.0807-0.43990.25250.1016-0.15850.29330.09780.00030.5925-0.16631.098823.138630.142191.2424
720.7160.62820.99692-0.97493.74680.08650.3927-0.268-0.2125-0.15160.4992-0.0444-0.30170.07160.38480.0307-0.00460.31220.03050.3915-1.291344.028979.7138
731.67821.915-2.87552.1864-3.28274.92730.0993-1.0346-0.63110.95120.04290.9630.6324-1.1915-0.14611.0195-0.05820.10231.0262-0.10481.221-18.0139-28.842377.9313
741.66770.2409-0.77350.0351-0.11350.3583-0.14270.1836-0.87980.1385-0.0160.83010.9358-1.06330.16830.4506-0.0787-0.0170.50540.08250.945-16.0617-21.914172.1839
754.11555.4830.40712.0105-1.89181.0471-0.06360.126-0.105-0.42150.27880.21050.1689-0.2929-0.12210.4495-0.0112-0.08210.3622-0.13531.002-10.3361-4.88164.5631
762.41623.805-1.32626.9596-1.58461.0001-0.13970.3457-0.148-0.49770.05210.05460.17520.02170.05030.46440.0762-0.01390.4103-0.14440.6759-3.84814.771558.0089
776.00680.54093.09181.99990.99364.8038-0.06280.3480.6746-0.51990.1053-0.3172-0.75320.2872-0.03810.57810.00320.15440.4492-0.17980.8399-2.659728.905760.2409
782.04080.45890.33223.314-2.33941.8660.10690.21060.9474-0.45880.1019-0.8046-0.85220.4605-0.20690.498-0.13810.16240.7752-0.12681.1039-0.240737.238865.9675
796.52197.8374-1.15852.0007-4.55443.43870.05410.2170.521-0.0417-0.0204-0.4737-0.64230.6762-0.03210.52460.00790.20370.5968-0.13730.88196.538623.192659.6312
802.02881.32121.46448.9443-2.13652.23890.07120.227-0.308-0.28420.0251-0.34290.40880.2962-0.03660.50470.0574-0.00260.313-0.16890.71743.11313.892664.843
812.00151.99353.08541.9983-2.25864.1668-0.4947-0.80260.410.88740.6471-0.1154-0.3621-0.1674-0.06980.37420.0297-0.03380.3256-0.04170.3939-8.743837.363274.2496
820.8156-2.8206-0.36029.77541.27590.1668-0.0005-0.0080.0438-0.10270.0877-0.69070.16360.2333-0.04040.3552-0.0014-0.02550.2719-0.01760.68494.529211.371273.7475
835.67083.28621.85242.00183.96175.9341-0.00120.34370.2308-0.690.0208-0.1691-0.17550.0841-0.0310.46210.12-0.07890.2741-0.00720.72816.6213-9.945270.9515
842.0016-4.2791-5.71584.16986.27939.50250.07860.51850.0087-0.5636-0.17480.0041-0.0839-0.21890.02870.39740.0601-0.17220.3280.0550.92264.3585-21.813365.6193
854.4614-4.838-3.63752.0031-3.6022.00110.2644-0.87930.32880.2797-0.29480.14980.0717-0.27040.07340.3373-0.1075-0.07510.47480.02010.97344.34-24.571373.8151
865.806-7.9563-2.78142.00032.29247.1908-0.0585-0.33840.03450.4673-0.03050.24940.0503-0.02030.05980.48040.0762-0.09420.31480.1020.87073.9067-13.591779.8048
875.3453-3.38440.71172.0023-6.37844.0925-0.0481-0.2270.11550.44710.0203-0.2381-0.08820.11170.00490.42870.0873-0.11970.3502-0.12590.6519-0.67411.534778.9376
883.7423-4.87822.19167.8658-3.31651.61730.0006-0.1011-0.1644-0.01440.14870.2478-0.075-0.0347-0.11390.24270.03180.01330.2526-0.06010.4502-15.621429.567171.8317
891.00880.86750.67560.73950.57010.43920.1257-0.2182-0.03740.2847-0.034-0.1799-0.00180.1467-0.0430.99170.1633-0.39181.24940.12161.170320.983740.3654134.7676
902.0008-6.5946-7.98063.86165.32429.4886-0.04-0.15070.2068-0.01490.06370.31870.054-0.12440.00820.72160.1178-0.37040.77430.10790.908413.938537.5852118.5648
910.9926-0.3303-0.05950.3753-0.00750.9158-0.1301-0.187-0.34180.32990.0812-0.01390.35490.04780.06040.50340.07560.06730.30690.04160.7088-5.816921.8991101.5136
922.7225-2.1680.13894.96270.79191.73890.16410.165-0.7377-0.2917-0.11860.41240.236-0.3308-0.08290.3982-0.04190.04310.4599-0.06350.7238-31.561823.690894.6666
934.6070.23480.21743.1270.39983.33520.1083-0.3429-0.40170.2367-0.13540.59110.441-0.98450.00550.4657-0.0324-0.04380.49380.02621.0668-16.305914.44778.6521
940.78230.1080.4428.84433.41124.72010.02130.1254-0.54160.0587-0.03520.22190.49980.050.02380.4927-0.0480.10560.4545-0.0290.7765-29.859419.234297.7527
950.68390.1348-0.21480.8641-0.00580.63250.01620.11670.299-0.2011-0.02630.2516-0.1512-0.3-0.04720.40080.05430.01310.38830.0070.4978-16.282846.149895.3742
960.7813-0.58920.29755.7733-1.03310.6246-0.0580.0789-0.4569-0.0460.24030.06040.0613-0.0669-0.06560.24180.02360.01930.3251-0.05540.6258-9.11097.411168.1596
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 59:72)
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 58:125)
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 58:69)
90X-RAY DIFFRACTION90(chain N and resid 70:76)
91X-RAY DIFFRACTION91(chain N and resid 77:179)
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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