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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 / homologyChromo/chromo shadow domain / Histone-fold / Inhibitor of growth proteins N-terminal histone-binding / Chromatin modification-related protein Yng2/Png1 / RNA binding activity-knot of a chromodomain / Enhancer of polycomb-like / Histone acetyltransferase subunit NuA4 / MOZ/SAS family / Core histone H2A/H2B/H3/H4 / MYST, zinc finger domain ...Chromo/chromo shadow domain / Histone-fold / Inhibitor of growth proteins N-terminal histone-binding / Chromatin modification-related protein Yng2/Png1 / RNA binding activity-knot of a chromodomain / Enhancer of polycomb-like / Histone acetyltransferase subunit NuA4 / MOZ/SAS family / Core histone H2A/H2B/H3/H4 / MYST, zinc finger domain / Zinc finger, PHD-type / Histone H2A / Histone acetyltransferase domain, MYST-type / Histone H2A/H2B/H3 / Zinc finger, FYVE/PHD-type / MYST family zinc finger domain / Zinc finger, RING/FYVE/PHD-type / Chromatin modification-related protein Eaf6 / Acyl-CoA N-acyltransferase / Chromo-like domain superfamily / Enhancer of polycomb-like, N-terminal / Zinc finger, PHD-type, conserved site / Zinc finger, PHD-finger / Inhibitor of growth protein, N-terminal histone-binding / Enhancer of polycomb protein / RNA binding activity-knot of a chromodomain / Histone H2A, C-terminal domain / Histone H2A conserved site / Winged helix-like DNA-binding domain superfamily / C-terminus of histone H2A / Histone acetyltransferase Esa1/KAT5/Tip60 / SIRT1 negatively regulates rRNA expression / Sensing of DNA Double Strand Breaks / Condensation of Prophase Chromosomes / MYST-type histone acetyltransferase (HAT) domain profile. / Zinc finger PHD-type profile. / DNA Damage/Telomere Stress Induced Senescence / Recruitment and ATM-mediated phosphorylation of repair and signaling proteins at DNA double strand breaks / Activated PKN1 stimulates transcription of AR (androgen receptor) regulated genes KLK2 and KLK3 / Senescence-Associated Secretory Phenotype (SASP) / HATs acetylate histones / Regulation of TP53 Activity through Acetylation / Zinc finger PHD-type signature. / RUNX1 regulates genes involved in megakaryocyte differentiation and platelet function / Estrogen-dependent gene expression / Histone H2A signature. / Oxidative Stress Induced Senescence / covalent chromatin modification / NuA3b histone acetyltransferase complex / NuA3a histone acetyltransferase complex / NuA3 histone acetyltransferase complex / Piccolo NuA4 histone acetyltransferase complex / histone H4-K12 acetylation / histone displacement / histone H3-K14 acetylation / histone H2A acetylation / DNA-templated transcription, elongation / H4 histone acetyltransferase activity / histone H3-K23 acetylation / nuclear-transcribed mRNA catabolic process, non-stop decay / peptidyl-lysine acetylation / NuA4 histone acetyltransferase complex / chromatin silencing at silent mating-type cassette / peptide N-acetyltransferase activity / histone H4-K8 acetylation / histone H4-K5 acetylation / nuclear euchromatin / positive regulation of macroautophagy / histone acetyltransferase complex / MOZ/MORF histone acetyltransferase complex / histone acetylation / chromatin silencing at rDNA / histone acetyltransferase / histone acetyltransferase activity / methylated histone binding / positive regulation of transcription elongation from RNA polymerase II promoter / regulation of cell cycle / meiotic cell cycle / nucleosome / chromatin DNA binding / chromatin organization / histone binding / chromatin remodeling / RNA polymerase II proximal promoter sequence-specific DNA binding / nuclear chromatin / cell cycle / regulation of transcription by RNA polymerase II / DNA repair / negative regulation of transcription, DNA-templated / chromatin binding / nucleolus / protein heterodimerization activity / positive regulation of transcription by RNA polymerase II / DNA binding / identical protein binding / metal ion binding / nucleus / cytosol / Chromatin modification-related protein YNG2 / Enhancer of polycomb-like protein 1
Function and homology information
Specimen sourceSaccharomyces cerevisiae (baker's yeast)
MethodX-RAY DIFFRACTION / SYNCHROTRON / MOLECULAR REPLACEMENT / 3.25 Å resolution
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.
Validation Report
SummaryFull reportAbout validation report
DateDeposition: Apr 11, 2016 / Release: Oct 26, 2016
RevisionDateData content typeGroupCategoryItemProviderType
1.0Oct 26, 2016Structure modelrepositoryInitial release
1.1Oct 18, 2017Structure modelAuthor supporting evidence / Database references / Derived calculationscitation / pdbx_audit_support / pdbx_struct_oper_list_citation.journal_id_CSD / _pdbx_audit_support.funding_organization / _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
L: Htz1
M: Htz1
O: Htz1


Theoretical massNumber of molelcules
Total (without water)318,34715
Polyers318,34715
Non-polymers00
Water0
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
Polyers106,1165
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area (Å2)19400
ΔGint (kcal/M)-108
Surface area (Å2)41400
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
Polyers106,1165
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area (Å2)18780
ΔGint (kcal/M)-108
Surface area (Å2)41640
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
Polyers106,1165
Non-polymers00
Water0
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area (Å2)18750
ΔGint (kcal/M)-111
Surface area (Å2)41000
MethodPISA
Unit cell
γ
α
β
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-MC 1 2 1

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Components

#1: Protein/peptide Histone acetyltransferase ESA1


Mass: 36556.883 Da / Num. of mol.: 3 / Fragment: UNP residues 141-445 / Mutation: E338Q
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/peptide Chromatin modification-related protein EAF6 / ESA1-associated factor 6


Mass: 12915.704 Da / Num. of mol.: 3
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/peptide Enhancer of polycomb-like protein 1


Mass: 41835.215 Da / Num. of mol.: 3 / Fragment: UNP residues 50-400
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/peptide 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: (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: (gene. exp.) Saccharomyces cerevisiae (baker's yeast)
Production host: Escherichia coli (E. coli) / References: UniProt: Q12692*PLUS

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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 / Density percent sol: 76.17 %
Crystal growTemp: 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 kelvins
SourceSource: SYNCHROTRON / Site: SSRF / Beamline: BL17U / Wavelength: 0.987 Å
DetectorType: ADSC QUANTUM 4r / Detector: CCD / Collection date: Jan 10, 2014
RadiationDiffraction protocol: MAD / Monochromatic or laue m l: M / Scattering type: x-ray
Radiation wavelengthWavelength: 0.987 Å / Relative weight: 1
ReflectionD resolution high: 3.25 Å / D resolution low: 30 Å / Number obs: 79960 / NetI over sigmaI: 6.9 / Redundancy: 3.4 % / Percent possible obs: 79.6

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Processing

Software
NameVersionClassification
PHENIX(1.10.1_2155: ???)refinement
HKL-2000data reduction
HKL-2000data scaling
autoSHARPphasing
RefineMethod to determine structure: MOLECULAR REPLACEMENT / Overall SU ML: 0.45 / Cross valid method: FREE R-VALUE / Sigma F: 1.45 / Overall phase error: 30.12
Solvent computationSolvent shrinkage radii: 0.9 Å / Solvent vdw probe radii: 1.11 Å
Least-squares processR factor R free: 0.2731 / R factor R work: 0.2502 / R factor obs: 0.2508 / Highest resolution: 3.25 Å / Lowest resolution: 29.996 Å / Number reflection R free: 1998 / Number reflection obs: 79960 / Percent reflection R free: 2.5 / Percent reflection obs: 78.86
Refine hist #LASTHighest resolution: 3.25 Å / Lowest resolution: 29.996 Å
Number of atoms included #LASTProtein: 19671 / Nucleic acid: 0 / Ligand: 0 / Solvent: 0 / Total: 19671
Refine LS restraints
Refine IDTypeDev idealNumber
X-RAY DIFFRACTIONf_bond_d0.00420081
X-RAY DIFFRACTIONf_angle_d0.62127039
X-RAY DIFFRACTIONf_dihedral_angle_d14.15012277
X-RAY DIFFRACTIONf_chiral_restr0.0422898
X-RAY DIFFRACTIONf_plane_restr0.0033485
Refine LS shell

Refine ID: X-RAY DIFFRACTION

Highest resolutionR factor R freeR factor R workLowest resolutionNumber reflection R freeNumber reflection R workPercent reflection obs
3.25010.39340.32093.331351197228.00
3.33130.35510.33243.421280316445.00
3.42120.36820.31973.5218106409358.00
3.52180.32600.30843.6353124487670.00
3.63530.33960.29953.7649146571081.00
3.76490.29480.28843.9154159618787.00
3.91540.30530.26944.0932160623189.00
4.09320.27370.26614.3084164639991.00
4.30840.23610.25114.5774164640591.00
4.57740.26600.23344.9294165644691.00
4.92940.26680.23255.4228170662694.00
5.42280.30830.24636.2015172672395.00
6.20150.25320.23527.7905171666494.00
7.79050.19360.181429.9975166646690.00
Refine TLS

Method: refined / Refine ID: X-RAY DIFFRACTION

IDL11L12L13L22L23L33S11S12S13S21S22S23S31S32S33T11T12T13T22T23T33Origin xOrigin yOrigin z
10.0064-0.01940.00960.01400.00210.03810.05550.1545-0.0225-0.06730.0365-0.03140.02710.0303-0.00000.2489-0.01410.03990.1294-0.05960.351310.830659.186875.4440
20.0321-0.02850.00950.0128-0.00510.0026-0.05180.06980.0278-0.08450.0559-0.10670.17280.10840.00000.0225-0.36320.2474-0.28240.05640.345911.545774.144670.5095
30.0031-0.00620.00060.01450.00520.01240.0902-0.11650.02490.01520.00980.03860.18620.18760.0000-0.5588-0.44890.4632-0.26950.3826-0.22785.505173.468584.0465
40.00190.00140.00160.00140.00000.0013-0.0040-0.0202-0.0065-0.0157-0.0322-0.0150-0.0080-0.01730.00000.1263-0.04710.11400.22450.07310.193519.142485.633774.7804
50.6094-0.2504-0.04210.20230.17390.2133-0.2603-0.2528-0.08260.1508-0.1322-0.06910.0851-0.04570.0002-0.0412-0.3482-0.2528-0.0710-0.23690.043113.276076.554594.8623
60.00220.00130.00170.00230.00210.00210.00730.0145-0.0016-0.0205-0.0144-0.0051-0.01350.0105-0.00000.4966-0.1968-0.10780.5194-0.23540.361618.186188.1059113.8670
70.01020.0092-0.01010.0042-0.00240.0076-0.03610.01000.0010-0.0230-0.00670.04010.0290-0.0062-0.00000.27440.0463-0.16720.53720.02260.149017.068277.7652113.8428
80.02070.0108-0.03870.0146-0.00280.03890.0228-0.01130.1616-0.03740.0373-0.0526-0.0512-0.01390.00000.0627-0.2247-0.18490.1929-0.16850.292615.005783.8332101.1445
9-0.00080.0004-0.00080.00060.0002-0.0003-0.0037-0.0029-0.00800.0022-0.0100-0.0025-0.0055-0.0060-0.00000.7679-0.1452-0.05570.62970.03410.648110.1735111.943427.3327
100.0039-0.00020.00690.01200.00000.0043-0.00920.03820.0156-0.00010.02400.0223-0.0041-0.00240.00000.3917-0.0965-0.16880.3872-0.05230.2058-2.3380112.910050.7283
110.00080.0001-0.00140.00230.00060.0015-0.00890.02580.00860.0284-0.02060.02350.0154-0.03050.00000.2072-0.0591-0.01390.2277-0.00020.2264-12.8485108.884172.9988
120.0007-0.0007-0.00130.00330.00010.00200.0062-0.00540.0032-0.0002-0.00220.00570.0004-0.0022-0.00000.58470.01710.01010.6435-0.01550.5311-16.4284100.962683.6559
130.00040.00090.00140.00070.00110.0015-0.01810.00610.01320.0078-0.0044-0.01020.0072-0.0003-0.00000.3127-0.03850.04990.23020.03270.1471-10.487396.869083.5123
140.00280.00100.00130.0056-0.00150.00380.01130.0057-0.0188-0.00030.01660.00930.01220.0125-0.00000.6268-0.0497-0.07640.5416-0.03010.5947-10.9643104.679385.6140
150.00050.0004-0.0007-0.0002-0.00190.0016-0.0181-0.0004-0.00550.01460.0290-0.00030.0185-0.00260.00000.2489-0.0554-0.02100.1690-0.05150.1349-2.4375109.155874.5964
160.0088-0.00110.00010.00080.00020.0004-0.00310.00140.0001-0.0031-0.00880.00130.00080.0035-0.00000.4243-0.0544-0.03950.45490.02660.47020.5687116.513871.6892
172.00525.82950.12776.19803.02882.97310.2560-1.0013-0.94870.43650.3643-0.46750.3854-0.2127-0.63080.00800.0836-0.23890.20340.07560.170237.96453.58689.639
180.00250.0003-0.00040.00040.00050.0001-0.01210.0157-0.0132-0.00750.00290.0004-0.0038-0.01000.0000-0.0563-0.0821-0.07500.10920.02150.048630.508664.461094.7302
190.03420.03790.01020.0139-0.0151-0.00530.07500.1235-0.0290-0.0977-0.0996-0.07950.05710.17400.00000.1538-0.16320.1982-0.03370.03800.360313.373273.948066.9925
200.0067-0.0000-0.00360.02150.01140.0021-0.0371-0.00420.0062-0.09870.05530.0976-0.05550.02550.00000.3283-0.06940.03190.20960.06880.3149-7.483266.491952.7181
21-0.00030.00600.00270.0053-0.00040.0083-0.02690.0196-0.0357-0.0100-0.05770.02040.0287-0.02510.00000.3185-0.0841-0.00160.2830-0.00070.2026-3.561589.980857.4799
220.0047-0.0102-0.00250.00750.01820.0164-0.04420.0322-0.0319-0.00940.09780.02930.03470.05040.00000.44710.02910.04010.23770.01190.2091-2.466965.549951.4016
23-0.0000-0.00590.00500.0059-0.00470.01090.04590.0137-0.0449-0.00500.0315-0.0516-0.0081-0.03690.00000.3636-0.05520.03990.1790-0.00410.1965-8.417561.558781.6971
240.06260.01660.1020-0.0299-0.05830.0144-0.29790.0210-0.2016-0.05890.11260.3480-0.18320.0793-0.00000.1603-0.1340-0.09770.2077-0.05980.1435-3.7597104.569658.2822
250.00120.0009-0.0009-0.00010.00020.0037-0.01000.0078-0.00560.01740.00680.00250.0295-0.00580.00000.2257-0.14020.03210.17320.17000.0830-18.234981.790681.6280
260.0044-0.00270.00210.0007-0.00050.00300.0204-0.0321-0.03170.00420.02590.02780.01920.02350.00000.22010.0256-0.01350.1830-0.10510.2066-3.581192.506278.4776
270.00190.00050.00210.00410.00720.00410.0087-0.0032-0.00750.00860.0144-0.0262-0.00600.0355-0.00000.3671-0.1997-0.14510.2916-0.09110.16079.2816107.322267.0562
280.0019-0.00030.00230.00210.00000.0051-0.02670.0258-0.0514-0.0081-0.00030.0082-0.0052-0.0006-0.00000.4570-0.2605-0.15120.39950.02170.168716.4070122.562447.3174
290.0084-0.00530.00810.00670.00680.0152-0.02350.0206-0.00440.01480.01190.00540.01710.0388-0.00000.2618-0.3126-0.03920.3790-0.03740.236717.4215109.598852.5319
300.00160.00180.00390.00180.00260.0079-0.04130.00580.00350.0109-0.0606-0.0095-0.00560.0383-0.00000.2764-0.2256-0.01190.2116-0.11910.1086-2.003294.118266.3457
310.0019-0.00080.00070.0033-0.00130.00350.00490.0258-0.0066-0.0026-0.01840.01040.00270.0251-0.00000.0943-0.2023-0.06540.34550.00630.2657-18.530584.790871.4654
320.00110.0006-0.0012-0.0003-0.00060.0009-0.01600.01090.0129-0.0091-0.00760.0144-0.0094-0.0079-0.00000.1057-0.08480.15560.20360.13080.2508-29.808383.707675.3688
330.0177-0.0049-0.01860.02290.00490.0167-0.0097-0.01410.0541-0.0387-0.00060.0558-0.00460.08880.00000.2000-0.0679-0.04210.2505-0.31720.2858-9.635581.1591109.3639
340.0009-0.00090.00000.0014-0.00070.00040.00140.00780.00740.0053-0.0133-0.0101-0.00850.01260.00000.43550.07570.00710.2188-0.03000.3685-11.718287.2670108.1109
35-0.02090.0097-0.01110.00070.0065-0.0086-0.0668-0.26190.0348-0.00610.0677-0.0557-0.0907-0.0886-0.00000.1554-0.0900-0.2280-0.0401-0.7130-0.6585-15.853470.4982119.2503
360.0008-0.00130.00070.00160.00030.00090.0040-0.0034-0.00950.00490.0034-0.0098-0.0109-0.0058-0.00000.6190-0.04740.00600.6273-0.09490.4872-15.421972.6527135.8079
370.1497-0.05920.10550.25570.04470.0594-0.0489-0.0755-0.1128-0.1161-0.1113-0.09110.03210.13270.0015-0.0053-0.1247-0.49790.2938-0.4570-0.6901-1.127357.7992123.5173
380.00330.0006-0.00280.00160.00700.0075-0.01800.0126-0.02350.00020.02470.04500.0087-0.01460.00000.35630.0636-0.36790.4970-0.05870.17578.141745.4654123.1179
390.00080.00010.00140.00020.00060.00060.00040.0065-0.0054-0.0004-0.0072-0.00120.0003-0.00220.00000.66760.0678-0.11610.7221-0.09020.725021.793744.7968124.4645
400.0010-0.01800.00110.04020.03330.04460.0044-0.1083-0.11220.0923-0.0342-0.03830.0196-0.02620.00000.33700.1935-0.32080.30370.0250-0.06680.824051.7904127.7334
41-0.0018-0.0009-0.00220.0008-0.00060.00020.00030.02800.0064-0.0099-0.00210.00870.01410.0175-0.00000.87690.12430.21250.93720.02790.7948-64.796289.0619152.9109
420.00010.0005-0.0010-0.0001-0.00010.0048-0.00350.00260.0069-0.00320.00090.0052-0.00740.0012-0.00000.8096-0.05530.08440.7959-0.01690.7528-62.263577.0508147.9954
430.00070.00010.0005-0.0001-0.00020.00110.01100.01100.0044-0.00630.0001-0.00070.00740.00020.00000.76520.01650.11410.8614-0.01750.6993-57.633767.4897144.4714
440.00000.0003-0.00100.0007-0.00080.00080.00170.0025-0.0106-0.0114-0.0104-0.00090.0159-0.0058-0.00000.6358-0.07260.09250.63150.04800.4788-52.136557.0538140.2708
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50-0.02120.0033-0.02510.00180.02240.0276-0.0593-0.08090.1684-0.0386-0.01140.0508-0.09620.04630.00000.3225-0.03040.04130.3387-0.4543-0.2932-24.156982.4973123.6972
510.0218-0.00020.00380.0173-0.01300.0132-0.0563-0.0346-0.0266-0.00770.07190.0047-0.0387-0.08170.00000.18190.01340.10150.5290-0.08190.4764-38.050786.6050105.4105
520.0047-0.0027-0.0007-0.00020.00230.00630.00570.01370.03170.05070.0474-0.0041-0.00200.0013-0.00000.2822-0.0779-0.00660.5870-0.09320.4279-43.420972.8859123.7732
530.01050.00050.01440.01540.00330.0110-0.08130.02530.0366-0.0505-0.0682-0.0277-0.0450-0.03810.00000.24100.14670.11230.4734-0.07880.3210-36.319490.1091108.3567
540.0202-0.01810.02010.01840.00020.0136-0.0281-0.0359-0.12670.05850.16300.09550.07040.0693-0.00000.1995-0.0267-0.03330.2683-0.13560.1529-26.110958.8661100.7174
550.03130.01220.00600.0178-0.01640.01100.0861-0.05640.02370.0834-0.00670.0570-0.0453-0.00330.00000.3105-0.09020.45860.5649-0.19150.1254-52.137271.3175145.7336
56-0.00000.0004-0.0022-0.0001-0.00050.00060.0146-0.00220.00470.01210.01160.0100-0.0092-0.00270.00000.82720.00290.05830.7980-0.06210.7370-56.631061.9564153.6860
570.0008-0.00100.0011-0.0001-0.00160.00200.0009-0.00820.0060-0.0061-0.00600.0037-0.00110.01080.00000.2696-0.07200.08360.3794-0.01070.2949-32.906051.8762104.4000
580.00450.00050.00240.00490.00140.00210.0036-0.0260-0.00720.03460.02840.0020-0.01520.0097-0.00000.3980-0.0718-0.01190.43000.01790.2787-30.486755.7455122.5684
590.00060.00050.00110.0006-0.00020.00100.0010-0.01780.00690.0001-0.01400.0023-0.00170.00170.00000.5907-0.0365-0.00870.6734-0.09190.4042-34.178860.4573141.1531
600.00050.00030.0003-0.00010.00060.00100.00550.0012-0.00230.0050-0.0062-0.0067-0.0010-0.00180.00000.64130.03170.07440.7227-0.05310.5820-39.349764.6744152.4697
610.00080.00040.0004-0.00130.00030.0004-0.00410.0124-0.0079-0.00710.0031-0.0010-0.0073-0.0050-0.00000.8017-0.03010.00830.9680-0.09800.7469-49.607469.8744161.8461
620.0001-0.00020.0004-0.00080.00050.0015-0.0077-0.0044-0.0026-0.0003-0.0177-0.00400.0025-0.0029-0.00001.25540.11080.06951.2861-0.04531.0319-50.783078.0439166.0973
630.01090.0110-0.00240.0103-0.00630.00250.0256-0.05020.0075-0.03980.03300.04830.02880.00410.00000.3809-0.0119-0.07640.5020-0.17280.1003-39.311769.5686139.9099
640.0060-0.0032-0.00600.00610.00240.0027-0.0462-0.0023-0.0021-0.0429-0.0507-0.03150.0004-0.00640.00000.3673-0.00040.05640.2764-0.03490.2753-41.747055.9177106.9371
650.00080.00200.00070.00020.00190.0024-0.02350.00920.00500.01980.01730.00680.0077-0.0087-0.00000.1526-0.0095-0.02240.19550.01840.1837-23.647341.9309105.1118
660.00400.00330.00220.0053-0.00110.0081-0.0153-0.01330.00560.0173-0.0067-0.00640.0298-0.0169-0.00000.23980.03960.06090.21600.05910.3262-5.363734.0434110.4327
670.0934-0.00430.05790.04140.04490.09310.2372-0.8297-0.07840.6339-0.1162-0.15950.48220.1561-0.0000-0.82491.1182-0.1277-1.02010.2325-0.0037-0.876130.049095.4658
680.15090.04200.00500.0978-0.11070.1553-0.0137-0.06980.0770-0.08570.1785-0.0738-0.16330.1591-0.0067-0.24320.40990.0249-0.04120.06570.443616.088141.548994.7492
690.00470.00080.00170.0113-0.00720.0047-0.01040.0046-0.0179-0.02520.0323-0.02110.0120-0.01070.00000.3179-0.0657-0.00970.23070.11360.191827.078051.469592.1637
700.0143-0.00500.01200.00450.00390.0075-0.0611-0.01940.02460.0126-0.0322-0.00350.06010.0662-0.00000.2575-0.1040-0.29520.36020.19350.549938.116149.428699.3743
710.0036-0.0014-0.00170.00240.00540.00110.0036-0.0161-0.0319-0.0409-0.0543-0.01950.01730.02990.00000.17330.0438-0.08020.3683-0.09250.305123.441429.904390.9524
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730.0005-0.00110.00000.00040.0002-0.00010.0037-0.0022-0.00030.01510.0053-0.00220.0033-0.0003-0.00000.45030.0202-0.02840.4528-0.08070.4741-17.8303-29.547877.7793
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760.0016-0.00090.00130.00170.00150.0013-0.02100.02040.00360.0010-0.02880.00530.03240.0362-0.00000.18870.0318-0.04850.1333-0.08980.2427-3.486814.232957.4804
770.0011-0.0020-0.00070.0001-0.00320.00270.00780.01830.0032-0.04190.02000.01520.0236-0.01320.00000.34030.04670.09710.3301-0.05880.3167-2.212728.479159.6728
780.00160.0002-0.00020.0004-0.00050.0002-0.0189-0.0094-0.0008-0.0144-0.00020.0160-0.01620.00110.00000.3302-0.05140.04990.3843-0.11180.47300.134136.808065.3773
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Refine 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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