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- PDB-8os6: Structure of a GFRA1/GDNF LICAM complex -

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Basic information

Entry
Database: PDB / ID: 8os6
TitleStructure of a GFRA1/GDNF LICAM complex
Components
  • GDNF family receptor alphaGFRα
  • Glial cell line-derived neurotrophic factor
KeywordsCELL ADHESION / Adhesion / Synapse / Complex / signalling
Function / homology
Function and homology information


RAF/MAP kinase cascade / : / diencephalon development / positive regulation of ureteric bud formation / postganglionic parasympathetic fiber development / positive regulation of monooxygenase activity / regulation of dopaminergic neuron differentiation / glial cell-derived neurotrophic factor receptor binding / regulation of morphogenesis of a branching structure / regulation of dopamine uptake involved in synaptic transmission ...RAF/MAP kinase cascade / : / diencephalon development / positive regulation of ureteric bud formation / postganglionic parasympathetic fiber development / positive regulation of monooxygenase activity / regulation of dopaminergic neuron differentiation / glial cell-derived neurotrophic factor receptor binding / regulation of morphogenesis of a branching structure / regulation of dopamine uptake involved in synaptic transmission / enteric nervous system development / positive regulation of branching involved in ureteric bud morphogenesis / peristalsis / sympathetic nervous system development / peripheral nervous system development / mRNA stabilization / metanephros development / neural crest cell migration / branching involved in ureteric bud morphogenesis / MAP kinase kinase kinase activity / growth factor activity / receptor tyrosine kinase binding / neuron projection development / signaling receptor activity / nervous system development / protein-containing complex assembly / negative regulation of neuron apoptotic process / receptor complex / external side of plasma membrane / protein homodimerization activity / positive regulation of transcription by RNA polymerase II / extracellular space / extracellular region
Similarity search - Function
: / Glial cell line-derived neurotrophic factor receptor, alpha 1 / Glial cell line-derived neurotrophic factor / Glial cell line-derived neurotrophic factor receptor, alpha 1/2 / Glial cell line-derived neurotrophic factor receptor / GDNF receptor alpha / Glial cell line-derived neurotrophic factor family / GDNF/GAS1 / GDNF/GAS1 domain / GDNF/GAS1 domain ...: / Glial cell line-derived neurotrophic factor receptor, alpha 1 / Glial cell line-derived neurotrophic factor / Glial cell line-derived neurotrophic factor receptor, alpha 1/2 / Glial cell line-derived neurotrophic factor receptor / GDNF receptor alpha / Glial cell line-derived neurotrophic factor family / GDNF/GAS1 / GDNF/GAS1 domain / GDNF/GAS1 domain / Transforming growth factor-beta, C-terminal / Transforming growth factor beta like domain / TGF-beta family profile. / Cystine-knot cytokine
Similarity search - Domain/homology
GDNF family receptor alpha / Glial cell line-derived neurotrophic factor
Similarity search - Component
Biological speciesDanio rerio (zebrafish)
MethodX-RAY DIFFRACTION / SYNCHROTRON / MOLECULAR REPLACEMENT / Resolution: 2.66 Å
AuthorsHoughton, F.M. / Adams, S.E. / Briggs, D.C. / McDonald, N.Q.
Funding support United Kingdom, 1items
OrganizationGrant numberCountry
The Francis Crick InstituteCC2068 United Kingdom
CitationJournal: Nat Commun / Year: 2023
Title: Architecture and regulation of a GDNF-GFRα1 synaptic adhesion assembly.
Authors: F M Houghton / S E Adams / A S Ríos / L Masino / A G Purkiss / D C Briggs / F Ledda / N Q McDonald /
Abstract: Glial-cell line derived neurotrophic factor (GDNF) bound to its co-receptor GFRα1 stimulates the RET receptor tyrosine kinase, promoting neuronal survival and neuroprotection. The GDNF-GFRα1 ...Glial-cell line derived neurotrophic factor (GDNF) bound to its co-receptor GFRα1 stimulates the RET receptor tyrosine kinase, promoting neuronal survival and neuroprotection. The GDNF-GFRα1 complex also supports synaptic cell adhesion independently of RET. Here, we describe the structure of a decameric GDNF-GFRα1 assembly determined by crystallography and electron microscopy, revealing two GFRα1 pentamers bridged by five GDNF dimers. We reconsitituted the assembly between adhering liposomes and used cryo-electron tomography to visualize how the complex fulfils its membrane adhesion function. The GFRα1:GFRα1 pentameric interface was further validated both in vitro by native PAGE and in cellulo by cell-clustering and dendritic spine assays. Finally, we provide biochemical and cell-based evidence that RET and heparan sulfate cooperate to prevent assembly of the adhesion complex by competing for the adhesion interface. Our results provide a mechanistic framework to understand GDNF-driven cell adhesion, its relationship to trophic signalling, and the central role played by GFRα1.
History
DepositionApr 18, 2023Deposition site: PDBE / Processing site: PDBE
Revision 1.0Nov 29, 2023Provider: repository / Type: Initial release
Revision 1.1Mar 27, 2024Group: Database references / Category: citation
Item: _citation.page_last / _citation.pdbx_database_id_PubMed / _citation.title

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Structure visualization

Structure viewerMolecule:
MolmilJmol/JSmol

Downloads & links

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Assembly

Deposited unit
A: GDNF family receptor alpha
B: Glial cell line-derived neurotrophic factor
C: GDNF family receptor alpha
D: Glial cell line-derived neurotrophic factor
E: GDNF family receptor alpha
F: Glial cell line-derived neurotrophic factor
G: GDNF family receptor alpha
H: Glial cell line-derived neurotrophic factor
I: GDNF family receptor alpha
J: Glial cell line-derived neurotrophic factor
K: GDNF family receptor alpha
L: Glial cell line-derived neurotrophic factor
M: GDNF family receptor alpha
N: Glial cell line-derived neurotrophic factor
O: GDNF family receptor alpha
P: Glial cell line-derived neurotrophic factor
Q: GDNF family receptor alpha
R: Glial cell line-derived neurotrophic factor
S: GDNF family receptor alpha
T: Glial cell line-derived neurotrophic factor
hetero molecules


Theoretical massNumber of molelcules
Total (without water)411,87038
Polymers405,88220
Non-polymers5,98918
Water34219
1


  • Idetical with deposited unit
  • defined by author
  • Evidence: electron microscopy, Negative stain electron microscopy reveals a particle with the same dimensions, 5-fold symmetry and architecture. Assembly also validated by Native PAGE and mutation of ...Evidence: electron microscopy, Negative stain electron microscopy reveals a particle with the same dimensions, 5-fold symmetry and architecture. Assembly also validated by Native PAGE and mutation of residues in the interface
TypeNameSymmetry operationNumber
identity operation1_555x,y,z1
Buried area53420 Å2
ΔGint-123 kcal/mol
Surface area139130 Å2
Unit cell
Length a, b, c (Å)114.140, 170.044, 130.791
Angle α, β, γ (deg.)90.000, 96.170, 90.000
Int Tables number4
Space group name H-MP1211
Space group name HallP2yb
Symmetry operation#1: x,y,z
#2: -x,y+1/2,-z

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Components

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Protein , 2 types, 20 molecules ACEGIKMOQSBDFHJLNPRT

#1: Protein
GDNF family receptor alpha / GFRα


Mass: 29249.277 Da / Num. of mol.: 10
Source method: isolated from a genetically manipulated source
Source: (gene. exp.) Danio rerio (zebrafish) / Gene: gfralpha1a, zgc:110129, gfra1a / Production host: Spodoptera frugiperda (fall armyworm) / References: UniProt: Q98TT9
#2: Protein
Glial cell line-derived neurotrophic factor / / zGDNF


Mass: 11338.884 Da / Num. of mol.: 10
Source method: isolated from a genetically manipulated source
Source: (gene. exp.) Danio rerio (zebrafish) / Gene: gdnf / Production host: Spodoptera frugiperda (fall armyworm) / References: UniProt: Q98TU0

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Sugars , 4 types, 18 molecules

#3: Polysaccharide
2-acetamido-2-deoxy-beta-D-glucopyranose-(1-4)-2-acetamido-2-deoxy-beta-D-glucopyranose


Type: oligosaccharide / Mass: 424.401 Da / Num. of mol.: 4
Source method: isolated from a genetically manipulated source
DescriptorTypeProgram
DGlcpNAcb1-4DGlcpNAcb1-ROHGlycam Condensed SequenceGMML 1.0
WURCS=2.0/1,2,1/[a2122h-1b_1-5_2*NCC/3=O]/1-1/a4-b1WURCSPDB2Glycan 1.1.0
[][D-1-deoxy-GlcpNAc]{[(4+1)][b-D-GlcpNAc]{}}LINUCSPDB-CARE
#4: Polysaccharide 2-acetamido-2-deoxy-beta-D-glucopyranose-(1-4)-[alpha-L-fucopyranose-(1-6)]2-acetamido-2-deoxy-beta- ...2-acetamido-2-deoxy-beta-D-glucopyranose-(1-4)-[alpha-L-fucopyranose-(1-6)]2-acetamido-2-deoxy-beta-D-glucopyranose


Type: oligosaccharide / Mass: 570.542 Da / Num. of mol.: 3 / Source method: obtained synthetically
DescriptorTypeProgram
DGlcpNAcb1-4[LFucpa1-6]DGlcpNAcb1-ROHGlycam Condensed SequenceGMML 1.0
WURCS=2.0/2,3,2/[a2122h-1b_1-5_2*NCC/3=O][a1221m-1a_1-5]/1-1-2/a4-b1_a6-c1WURCSPDB2Glycan 1.1.0
[][D-1-deoxy-GlcpNAc]{[(4+1)][b-D-GlcpNAc]{}[(6+1)][a-L-Fucp]{}}LINUCSPDB-CARE
#5: Polysaccharide alpha-L-fucopyranose-(1-6)-2-acetamido-2-deoxy-beta-D-glucopyranose


Type: oligosaccharide / Mass: 367.349 Da / Num. of mol.: 1 / Source method: isolated from a natural source
DescriptorTypeProgram
LFucpa1-6DGlcpNAcb1-ROHGlycam Condensed SequenceGMML 1.0
WURCS=2.0/2,2,1/[a2122h-1b_1-5_2*NCC/3=O][a1221m-1a_1-5]/1-2/a6-b1WURCSPDB2Glycan 1.1.0
[][D-1-deoxy-GlcpNAc]{[(6+1)][a-L-Fucp]{}}LINUCSPDB-CARE
#6: Sugar
ChemComp-NAG / 2-acetamido-2-deoxy-beta-D-glucopyranose / N-acetyl-beta-D-glucosamine / 2-acetamido-2-deoxy-beta-D-glucose / 2-acetamido-2-deoxy-D-glucose / 2-acetamido-2-deoxy-glucose / N-ACETYL-D-GLUCOSAMINE / N-Acetylglucosamine


Type: D-saccharide, beta linking / Mass: 221.208 Da / Num. of mol.: 10 / Source method: isolated from a natural source / Formula: C8H15NO6
IdentifierTypeProgram
DGlcpNAcbCONDENSED IUPAC CARBOHYDRATE SYMBOLGMML 1.0
N-acetyl-b-D-glucopyranosamineCOMMON NAMEGMML 1.0
b-D-GlcpNAcIUPAC CARBOHYDRATE SYMBOLPDB-CARE 1.0
GlcNAcSNFG CARBOHYDRATE SYMBOLGMML 1.0

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Non-polymers , 1 types, 19 molecules

#7: Water ChemComp-HOH / water / Water


Mass: 18.015 Da / Num. of mol.: 19 / Source method: isolated from a natural source / Formula: H2O

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Details

Has ligand of interestN

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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: 3.67 Å3/Da / Density % sol: 66.48 %
Crystal growTemperature: 303 K / Method: vapor diffusion, sitting drop
Details: 100mM Tris pH 7.5 100mM NaCl 3% (v/v) Acetonitrile 5% (w/v)PEG 20k
Temp details: Room Temperature

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Data collection

DiffractionMean temperature: 100 K / Serial crystal experiment: N
Diffraction sourceSource: SYNCHROTRON / Site: Diamond / Beamline: I04-1 / Wavelength: 0.92 Å
DetectorType: DECTRIS PILATUS 6M-F / Detector: PIXEL / Date: Apr 30, 2017
RadiationMonochromator: Single Bounce / Protocol: SINGLE WAVELENGTH / Monochromatic (M) / Laue (L): M / Scattering type: x-ray
Radiation wavelengthWavelength: 0.92 Å / Relative weight: 1
ReflectionResolution: 2.65→90.45 Å / Num. obs: 140996 / % possible obs: 99.1 % / Redundancy: 13.5 % / Biso Wilson estimate: 77.01 Å2 / CC1/2: 0.99 / Net I/σ(I): 4.75
Reflection shellResolution: 2.65→2.7 Å / Redundancy: 12 % / Mean I/σ(I) obs: 0.25 / Num. unique obs: 6703 / CC1/2: 0.281 / % possible all: 95.5

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Processing

Software
NameVersionClassification
PHENIX1.20.1_4487refinement
DIALS2.2.5data reduction
DIALS2.2.5data scaling
PHASERphasing
RefinementMethod to determine structure: MOLECULAR REPLACEMENT
Starting model: 3FUB
Resolution: 2.66→90.45 Å / SU ML: 0.529 / Cross valid method: FREE R-VALUE / σ(F): 1.33 / Phase error: 37.4716
Stereochemistry target values: GeoStd + Monomer Library + CDL v1.2
RfactorNum. reflection% reflectionSelection details
Rfree0.2828 6139 4.9 %Random
Rwork0.2437 119036 --
obs0.2456 125175 88.12 %-
Solvent computationShrinkage radii: 0.9 Å / VDW probe radii: 1.11 Å / Solvent model: FLAT BULK SOLVENT MODEL
Displacement parametersBiso mean: 82.95 Å2
Refinement stepCycle: LAST / Resolution: 2.66→90.45 Å
ProteinNucleic acidLigandSolventTotal
Num. atoms23416 0 390 19 23825
Refine LS restraints
Refine-IDTypeDev idealNumber
X-RAY DIFFRACTIONf_bond_d0.002424243
X-RAY DIFFRACTIONf_angle_d0.46932720
X-RAY DIFFRACTIONf_chiral_restr0.04013694
X-RAY DIFFRACTIONf_plane_restr0.00364245
X-RAY DIFFRACTIONf_dihedral_angle_d12.78199185
LS refinement shell
Resolution (Å)Rfactor RfreeNum. reflection RfreeRfactor RworkNum. reflection RworkRefine-ID% reflection obs (%)
2.66-2.690.6975410.6183805X-RAY DIFFRACTION18.09
2.69-2.720.5193620.52171280X-RAY DIFFRACTION28.35
2.72-2.760.4248920.46541841X-RAY DIFFRACTION41.16
2.76-2.790.51141040.47591907X-RAY DIFFRACTION42.53
2.79-2.830.42111350.42922901X-RAY DIFFRACTION64.49
2.83-2.870.46171910.43343488X-RAY DIFFRACTION77.58
2.87-2.910.43742320.4224192X-RAY DIFFRACTION94.25
2.91-2.950.39251940.40964310X-RAY DIFFRACTION95.38
2.95-30.41672340.41324429X-RAY DIFFRACTION98.33
3-3.040.46082140.41724457X-RAY DIFFRACTION98.67
3.04-3.10.40012350.38924429X-RAY DIFFRACTION98.75
3.1-3.150.37892320.35724446X-RAY DIFFRACTION98.55
3.15-3.210.36632440.34624377X-RAY DIFFRACTION98.74
3.21-3.280.33921990.33164457X-RAY DIFFRACTION98.62
3.28-3.350.34592260.32034502X-RAY DIFFRACTION99.02
3.35-3.430.32712430.30364408X-RAY DIFFRACTION99.04
3.43-3.520.35232270.29364449X-RAY DIFFRACTION99
3.52-3.610.29682320.28964439X-RAY DIFFRACTION99.24
3.61-3.720.32862290.28354496X-RAY DIFFRACTION99.16
3.72-3.840.29632260.26634463X-RAY DIFFRACTION99.28
3.84-3.970.27392360.23174461X-RAY DIFFRACTION99.24
3.97-4.130.24992210.21614463X-RAY DIFFRACTION99.3
4.13-4.320.26532300.20074481X-RAY DIFFRACTION99.37
4.32-4.550.23812300.19364532X-RAY DIFFRACTION99.56
4.55-4.830.20472380.18554450X-RAY DIFFRACTION99.51
4.83-5.210.21232190.17744532X-RAY DIFFRACTION99.56
5.21-5.730.23532510.18714466X-RAY DIFFRACTION99.6
5.73-6.560.28052510.21714501X-RAY DIFFRACTION99.58
6.56-8.260.24522300.19844528X-RAY DIFFRACTION99.62
8.26-90.450.23492410.17514546X-RAY DIFFRACTION98.68
Refinement TLS params.

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

IDL112)L122)L132)L222)L232)L332)S11 (Å °)S12 (Å °)S13 (Å °)S21 (Å °)S22 (Å °)S23 (Å °)S31 (Å °)S32 (Å °)S33 (Å °)T112)T122)T132)T222)T232)T332)Origin x (Å)Origin y (Å)Origin z (Å)
15.17087445516-3.13016690916-0.9091995396620.9063764268071.820034794865.11186721689-0.3426360511-0.2217809486150.181839428552-0.07627378313830.269334551713-0.5463420628170.230910339351-0.1712700827170.07745012862630.724218808473-0.0211881273035-0.03503372848330.242433378373-0.001044051259310.641617680057-5.7131.81281.67
25.786045883180.3804176367623.145409666171.464029732710.2307237983491.40478816005-0.1095110016020.0227319441755-0.799965566669-0.5028997675830.2872700193660.07425959294170.2745412906230.126109765464-0.1458711499210.713263109481-0.02692439867320.03200516014540.3824927092740.02182630930580.7623891341695.462-3.72989.979
33.45314770727-0.265681996817-2.385386403011.500409161740.6038709672724.70786374565-0.0932465826632-0.220906108625-0.0292886832920.1300557625730.234188916982-0.1953483891680.287610793350.196903151135-0.1550901076770.7996877776410.0377026172885-0.06793665349490.274723465821-0.002022196411730.6742365662692.1284.67299.712
43.083777215122.1196390539-2.665741427713.15840684156-1.97253777519.227853940280.1592687659640.3580508215320.0909077848008-0.01791415109970.3227661315480.408635297991-0.150985850593-1.22963983188-0.4540363969250.6656135381080.00309618060145-0.02873102024720.665967983551-0.0718611837570.657598264487-21.3542.49263.034
57.95712386656-2.258766721383.445591788195.0961128034-0.961260435532.77711497119-1.27780127741-0.124331524991-0.9554007284480.217527861161.469818922631.004406424571.10532679482-1.98748836053-0.07029758874151.08328731667-0.279989669610.2047299158951.19273024970.2090697673770.776407960026-23.6981.03133.893
64.20688185917-1.62537138601-0.3539968707768.09988707562-4.842587996382.69979586620.19370216319-0.006992289757960.30577245735-0.839633115072-0.09317526815-0.6177532607060.0769704900944-0.494228016299-0.1766309467220.661450627542-0.07198911136070.03953646409110.471963411712-0.02918640832140.631579403794-12.2270.57665.244
73.585516113170.505694644416-0.5059716930271.414179923340.3273538390282.808664113290.2006317279820.189875025157-0.229120943993-0.1379269059730.25617152525-0.7323137563770.30240361007-0.225324406226-0.4264649879180.9144235083880.127210075764-0.07678221658690.4118310212290.07170165586720.77393448607-12.6238.98176.927
83.18541202851-1.05957263584-2.573839056264.81302642776-2.883449566479.677123294080.806107433168-0.328844137395-0.5129115442470.759375168158-0.239200805462-0.00719376954913-2.00477571457-0.371579156637-0.5410270218460.9398044189180.033946357591-0.1400253512840.368386858992-0.0782792093640.781071003376-14.5645.88390.124
95.394841417651.732648112410.6891890510542.60069230851-2.876975739034.91686996978-0.7069735504440.3521547392570.3694608663490.2729482211890.3306586441920.977912748636-0.30550093061-0.886802499420.2782010771410.5651782399730.0321515891745-0.1354479054750.448499312055-0.01428700405510.730756970296-20.35238.69185.283
106.51976703731-2.698347188811.80945004251.50114460232-1.625796651511.79896001305-0.334479577590.412474490181-0.3737247445910.4561104342890.09970348501750.410463435346-0.09490602148270.006886250201880.2255711414040.734664266880.1472170695570.00845156294770.3778345653570.02798225038790.803049800441-12.99120.53489.223
112.334361984970.5313228395551.959467509093.085921333380.6395880857644.25606091465-0.6760335392560.574025645830.429132959594-1.191127605860.277113912888-0.1995276076520.4485673969481.493336718690.4189457250781.08183339132-0.03822808026940.1812429089110.5241695308170.1428998343560.733372842998-0.93528.28193.339
121.953436216531.63537747593-0.6156141135931.237236182-0.01748308735717.008004631190.510888464648-0.09365433902930.1690157152080.328717529880.007311280483240.1820705827770.7132784358481.00477939028-0.5755510034630.6259294471830.1144577296930.02544788851340.35453634238-0.02574646275650.639795568053-4.22638.016102.382
133.80606858724-2.48065094347-0.4664486667765.859062449352.367823746285.69147075739-0.016828266483-0.110554923353-0.2772924430020.777116897386-0.9396541396130.3734825391971.52707201059-1.791242106620.487350468130.9002991162140.3042485498970.009682915632830.1717832297510.1935824249780.617300038556-16.63933.16192.764
142.10754483309-1.209985894982.444832434272.026003528884.246773846632.395631381950.541381423065-1.791007336530.28747584719-0.782688655614-0.288035496667-0.1665990986740.452596817887-0.469460580316-0.217467534270.643817524172-0.09041542543540.06458265559370.7571510191130.07755003964970.929443579767-17.99417.01798.7
157.072355315780.620654261114-0.1913618761088.99212591211.366859762954.37838838497-0.31611423829-0.09102070905530.3445601876560.7673570968390.7721459056791.36492500307-0.284434827901-0.478079902811-0.5005197182370.5607681606460.03519719232220.1043058247170.505328641650.06274966363650.411669784391-12.17135.407100.625
165.609159738761.08813370569-3.55725542520.312940172732-1.645551378438.02627930785-0.872303032228-0.2987398816390.2144055064890.1468292547830.133204343401-0.4392219261840.1889592948730.3065889719340.5599747300760.8712096361980.0675936585907-0.002510492185150.387912075484-0.06616884029840.78283356696-9.48661.90662.258
172.067766498950.722541235171-1.301450213437.412381542931.010242804735.88160022003-0.277943546679-0.4614772979450.1130882412620.5690756405630.7607956369570.8782882469050.381217657833-0.582095863117-0.4957464047170.8431411952380.254180835865-0.008060441868690.5292325949480.1349142092850.868811577295-26.2651.95171.779
180.8465681829860.185468592889-1.2417195006-0.06738414919590.3378491302563.931080868540.68595496034-0.01408459935850.393879967295-0.4267535963560.3912939300680.472372249287-0.1375303698420.40848053077-0.838735396711.09501101140.0951854785262-0.0389938197670.312627157279-0.03414501836320.861718275744-10.76561.21456.912
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703.712024481160.8932909611662.848395821516.88198270566-1.720840263274.871293007380.1345889695110.510764172928-1.130488355690.4600187135140.0724479787313-0.08387895106462.164956298490.428651351753-0.1899766162351.3204144170.1041090596420.1775315568690.578409993724-0.1025564226010.85814804224533.738-24.39564.955
712.847818971860.6315937188782.950575820261.608483594561.267858757076.801798804980.2544096921710.139680064817-0.0525625208139-0.344011856262-0.0206771070369-0.06947493350980.1782026842970.246615025415-0.269110696731.0274129580.2151294142150.01818099768970.4743502262260.07269087225690.74915523073827.456-14.00333.843
722.84510387705-2.744769698780.07424102674172.476573385040.3769693190863.83194666520.4821991216220.182385296764-0.105165925507-0.543424990219-0.393185729788-0.254599736803-0.820523318789-0.2183217975540.007443661723570.967832314313-0.038402288459-0.01270181694910.370964760991-0.06479732899560.82296481040241.54459.61222.679
734.89872039745-5.243185795185.852663581478.37601029585-4.113255026942.033120068520.3766613327260.849631486758-0.5292547363980.389524624568-0.940185960015-0.4977190300720.5275350579761.483388981130.4561257648111.021818191760.015152146726-0.0168599371360.9329875774950.1217742758450.7261004799856.82445.50130.573
746.11963427641-3.305217890621.616083165146.17735817173-0.674780458997.087433709361.0215896417-0.528667877393-0.801731328828-0.476556218462-0.3104192892660.5854523847051.29769808178-0.511967187096-0.6496713141630.882393464826-0.0863233994178-0.07822937311570.581005916708-0.06700108058250.79389231993442.49945.2223.278
754.90375506683-2.19468358215-4.12596254222.411136424272.110162994378.871436323690.3341552717390.684071136227-0.77042645802-0.477985324318-0.612505453119-0.109480750193-1.90616862260.06447790057650.4614344303690.9727663255270.0381338891461-0.08924873351790.630001658656-0.03842240844730.71104693353543.50258.03811.336
764.90164199791.98090796179-0.6603080427868.552478712712.98728277613.91990764959-0.107932911093-0.351661521393-0.438059886083-0.8618822752670.0628785165945-0.07007481106370.5399519459331.089883113910.09181432958060.7759880677220.131469346395-0.03329013051860.5626633478770.07148345292260.55468804816353.67643.64614.593
776.538822903070.3151617078180.8572860346147.55255862404-2.174190549796.105987374140.160542649681-0.949629441048-0.242325443361-0.118494477520.01086355172260.6725629789120.0733046721686-0.761446127048-0.134482635350.787424942930.09476453695660.1372826931130.7411769119250.05922759105460.73333033145940.54635.9628.878
780.4191570443590.314571577507-1.219154641881.6426981676-0.7472558851997.199727699780.3122735532240.2165381576080.0729677395362-0.1456932761370.211997029508-0.1933511918521.07092290434-0.659277987762-0.5206210642720.705485807559-0.01634465047560.006007561855580.5989931654020.02383292175430.72654854417538.90745.767-0.595
796.61025960765-1.996474756953.094744546482.29995977562-6.219455906075.580707184070.2352719353890.03833963890360.417432456324-0.830472351474-1.33131373399-1.125297392340.1924100151431.07154197371.01175022130.7456336528210.1394581785990.0002283252538120.703098661115-0.0603929424660.6874962534451.78848.0119.182
808.11801313862-0.06834802517634.279778084154.87835830117-1.43659904169.392694538490.08683829100161.14178929778-0.3864433444150.015454226119-0.304461501287-0.5411117933140.1432017675741.127523503190.1786000534270.749759634687-0.1657028231120.04729131703670.715234517764-0.06275543206380.76076384392356.01937.2081.293
814.98306889455-2.965676511912.766120641842.20492253727.324642434538.73602716454-0.275163123242-0.08224916239850.607274125114-1.997284557070.700932171939-0.383490785822-1.233861698250.474871648884-0.2968844366790.767433001714-0.07618667528710.03810400353030.6103498054130.01123840794810.60459900504541.39555.812.605
824.19212259001-1.03490021056-0.1250935860513.45140029915-0.7148504735193.510083570390.234205977975-0.4935345452270.0492318520607-0.0558570486022-0.201718863062-0.376120446057-1.072797688210.228509240540.00970385576431.02898977591-0.1487876602470.01692363591950.6392182580380.04307618575950.64438227114943.5871.39831.274
832.53931245356-0.83424712159-3.375002592411.299275011430.24458505301710.09999144550.910031773980.2482608027010.2717684060610.0588160957274-0.09366088662670.0285563952076-1.70926229353-1.07621345922-0.9454451462321.099311597850.03393309291180.000356533413250.5541900522330.04364950353010.84665031398633.05972.53343.506
845.7047398327-3.909838516510.8954326257013.161696343531.706488651799.522564393960.9316100973490.4637374326421.361262817670.110829609455-0.258689441489-0.810047405457-1.72643109830.454354984366-0.6679388593070.935417066574-0.135774631550.008914788591210.555027436571-0.02814806481250.85245628468433.72377.88166.325
852.970849857841.564308994082.390744727922.771401906093.057755943383.489398365170.3171674485420.205512671940.632093198750.02742196587760.000470047189543-0.745253509976-1.826774557861.31191263873-0.176032699650.824243538208-0.3868814112340.09863757668740.822746928889-0.01483896779981.0124963227140.78476.92964.061
861.18778586888-1.5474543314-0.772570905712.718405963431.45051816873.404150521420.464939440568-0.120455500550.168928422148-0.688732730622-0.0408442888176-0.3858073491110.0774213335578-0.00367208978923-0.4193456156560.879391818726-0.07335494808930.002202277546450.389013676692-0.0103701808680.70958610971142.75962.69135.572
874.129678960732.43252587001-0.023942554325.256132335611.362629307899.63331526785-0.1241994460820.0178774250849-0.403732401862-0.2985307563780.00724706777081-0.09886515243360.2282530026541.219274789510.1454015993320.5897513113050.105611323782-0.01836527157280.6306842800230.07514972001850.60636313726857.57214.70121.439
882.27270470490.5196308895670.05492113856210.4562670499271.499581009882.2736120743-0.141659349673-0.162288548024-0.1279846256550.1765035556430.1244822433480.0298357246894-0.065868508752-0.09310608660380.04901351351610.7994640486930.02179063025380.01754521844850.5186625579760.01271441093970.69267503147745.51612.8717.622
896.88503382882.25402879548-5.020428537052.62772437240.2941751774465.01223939619-0.816379176921-0.38261815632-1.10537069837-0.8217706701360.0259458787639-0.2906867726071.041445512750.4372709350750.6138234618170.860519839904-0.02781806076450.02180122048470.628247695277-0.02259100322210.61776715497153.958.1919.485
906.55695165653.31741211166-0.6688256337927.16646954267-2.401538254845.66660432674-0.1761435680650.217194290841-0.132909457731-0.3297656303890.269898199760.146836669536-0.150261097113-0.137119506809-0.09170010184120.7957824632320.231833659606-0.01632779724530.6089244971950.02001504009060.62789738271350.4989.4151.74
916.219006570181.242161366940.9566529179097.342580149544.884952515832.2843817703-0.132488418524-0.06948680648360.09925913004440.3223483451370.554154483911-0.510459611630.677835782252.33683536994-0.465614339370.8132541536930.07969495416340.03221901788021.208747151290.1133759749370.61787145649671.54521.63935.49
922.840924470163.17370856484-0.9977109756079.607454567914.280438105775.55118001212-0.02634631090490.2313897152730.360627926823-0.9688681523780.359732580532-0.922113024609-0.870567440692.43220533751-0.3759126368030.632890104873-0.1675414960880.03789169084421.251766008730.04640473324420.74957161136871.27930.8166.268
930.8882690325750.9408517053860.436066200424.018811611550.6746255110070.2143452442440.1140592594740.426545573881-0.0505011107407-0.731253317367-0.762023654112-1.462215317530.8515154635080.2811664095030.6074736244310.7387520067620.1056339653250.1616698798671.18564743794-0.05707547837851.1864673772774.48321.64167.807
942.37932389143-2.10530666061-2.082459695054.425531591062.853996349116.159509733710.2275239763050.100416853645-0.179812141108-0.1874312422630.0479818910591-0.0323420629733-0.3935489044280.324400869005-0.3147241475680.5641815162310.0577309350863-0.02722306821091.034606399590.07550447318750.81144593080263.49718.05237.72
956.659607172510.3020787743793.282301046348.093056370950.693921804126.585520008160.130646667938-0.1157424177680.40941846632-0.1291574174530.132189705956-0.1360276201941.09000741182-1.01289247343-0.2800323317870.702119339021-0.07001071716230.1084260661170.6090968005440.007219454841830.474518573749-9.31912.09117.523
962.691211664642.04986462997-1.947898027056.056706881181.965505587878.628797829720.112119675159-0.05903579464341.054589149020.1131379614630.0130419621178-0.314181478873-1.13702842942-0.787020401014-0.3241109471640.7800320048290.09898068224710.08760360537390.533586936493-0.06015335016320.633530554379-6.28927.11420.205
973.53032783249-0.84477505602-0.5314408170876.60086755063-0.3624563887645.876059140970.102254774550.0746959208620.0916643735032-0.728521181788-0.0481185767033-0.251280259191.01546596222-0.682830409638-0.07429761984190.896040995797-0.04022877116410.1551144051370.4857990979930.01796802772960.603517538042-10.41616.0146.95
984.262675688015.973696935590.3405849720868.102631592760.5403010103125.361212676830.1752692368690.02745550215960.1493358334420.02798414651530.2532622283471.33661878661-0.84972637623-0.220949005787-0.3288275228220.967699818530.08854278727110.07368934992380.413132976628-0.00538355050070.904110984223-3.15237.5726.922
995.55237522856-2.25350227138-5.100095220696.684624669784.372527661666.604449486530.33266903558-1.94011203667-1.530820619171.65339864294-0.005990764538340.3094643677910.3727727384250.989735362927-0.4283439248310.5373716179170.305986053657-0.07425270563240.389750553079-0.01640443534551.018224637718.34128.0735.422
1000.586531378261-0.600213539434-1.330866562321.9634473505-0.1111232335885.02440089188-0.02390676020890.0456510090031-0.126639110149-0.797044260762-0.1264932850150.346071916697-0.1636169588010.4517403943270.05893920058760.7018211528850.048423803425-0.008174528172980.348168995124-0.02816974640910.5907843725735.81718.668-4.182
1016.226357054972.50736053641.305481361254.4555481335-3.345522389665.526652772280.6144331253990.2248502105910.4544524003960.0824839843158-0.6814831595260.7474843133120.47083119242-0.04004665552450.02599964905340.9351827641870.08934714871270.08356644664480.3675337088760.01114719278520.584091402157-7.73324.9012.831
1027.67261258371-6.114801840682.851695774798.095078482020.6617880053475.72526993011-0.0869256653231.15349019651.83249532858-1.00530406228-0.74909892707-1.74299694992-2.086937352680.9556802220050.603299238429-0.025705752180.899134304307-0.09672607452070.7739832453460.4008350838081.20145615187-6.87736.669-0.692
1032.25426376804-0.00138591115591.785586989539.496147451980.735466000632.189815187690.2549114360350.207757625511.08943003246-0.909338594101-0.4353881590290.677373465774-0.885433412348-0.6634598016540.08751915677820.8799498688350.00736289642286-0.02969115606820.737882199701-0.008694622567290.672495689836-0.90335.806-6.254
1042.133880847772.3582595478-2.445043389535.812868630455.068976853686.92372248838-0.3061488528290.1920368712020.349591136932-0.4165304396710.6334246684241.37252123170.3138929424580.158182145167-0.2129661814830.5890976725120.00149949568039-0.05223279497440.4556419563790.03066828273880.574805667556-2.4914.291-3.05
1052.280956118131.216527824621.903561753454.68772727111-1.868962056032.677061566670.21078853655-0.06252662320350.0304599324562-0.5889810888550.5431799205660.2989245481761.77956132556-1.34425901466-0.7505695379061.03707263999-0.08359905650590.1116241398720.8710117262140.08021622540340.725278222836-17.1090.85930.427
1067.07646435107-3.03395167821-0.1544423498362.21697897453-0.5489341623312.062257588040.4732178271520.376438713566-1.70654367489-0.89514603412-0.40203085012-0.7673266568672.12227167806-2.44927532269-0.1496438168141.1300875387-0.307026472060.02755488347090.68240694537-0.1271332270210.791750636924-21.388-5.60760.277
1075.484697938226.04330776723-3.2236955892.14546116122-6.805816384057.08059272215-1.22787988109-0.5343018215920.747003782506-0.9350671697080.18267265401-0.03597491702240.282227457812-0.5824283293520.9257412677340.911838897243-0.109348392627-0.1634405738011.0515844824-0.0484454133230.74515535097-26.366-0.90356.853
1084.78329640185-1.798092300644.474280921667.42137362526-4.680570444037.17967321292-0.1818574855810.1526000163480.05694795178810.5898919102170.270908346327-0.132334742528-0.6766014570450.0684622447667-0.09763288494490.715203193824-0.1188224140910.1282170112460.683315424537-0.07150855058110.554137125592-14.40210.22729.307
Refinement TLS group
IDRefine-IDRefine TLS-IDSelection detailsAuth asym-IDAuth seq-ID
1X-RAY DIFFRACTION1( CHAIN A AND RESID 149:222 )A149 - 222
2X-RAY DIFFRACTION2( CHAIN A AND RESID 223:256 )A223 - 256
3X-RAY DIFFRACTION3( CHAIN A AND RESID 257:355 )A257 - 355
4X-RAY DIFFRACTION4( CHAIN B AND RESID 139:178 )B139 - 178
5X-RAY DIFFRACTION5( CHAIN B AND RESID 179:201 )B179 - 201
6X-RAY DIFFRACTION6( CHAIN B AND RESID 202:235 )B202 - 235
7X-RAY DIFFRACTION7( CHAIN C AND RESID 151:202 )C151 - 202
8X-RAY DIFFRACTION8( CHAIN C AND RESID 203:222 )C203 - 222
9X-RAY DIFFRACTION9( CHAIN C AND RESID 223:237 )C223 - 237
10X-RAY DIFFRACTION10( CHAIN C AND RESID 238:256 )C238 - 256
11X-RAY DIFFRACTION11( CHAIN C AND RESID 257:269 )C257 - 269
12X-RAY DIFFRACTION12( CHAIN C AND RESID 270:295 )C270 - 295
13X-RAY DIFFRACTION13( CHAIN C AND RESID 296:314 )C296 - 314
14X-RAY DIFFRACTION14( CHAIN C AND RESID 315:325 )C315 - 325
15X-RAY DIFFRACTION15( CHAIN C AND RESID 326:354 )C326 - 354
16X-RAY DIFFRACTION16( CHAIN D AND RESID 138:150 )D138 - 150
17X-RAY DIFFRACTION17( CHAIN D AND RESID 151:162 )D151 - 162
18X-RAY DIFFRACTION18( CHAIN D AND RESID 163:178 )D163 - 178
19X-RAY DIFFRACTION19( CHAIN D AND RESID 179:190 )D179 - 190
20X-RAY DIFFRACTION20( CHAIN D AND RESID 191:207 )D191 - 207
21X-RAY DIFFRACTION21( CHAIN D AND RESID 208:224 )D208 - 224
22X-RAY DIFFRACTION22( CHAIN D AND RESID 225:235 )D225 - 235
23X-RAY DIFFRACTION23( CHAIN E AND RESID 150:212 )E150 - 212
24X-RAY DIFFRACTION24( CHAIN E AND RESID 213:222 )E213 - 222
25X-RAY DIFFRACTION25( CHAIN E AND RESID 223:237 )E223 - 237
26X-RAY DIFFRACTION26( CHAIN E AND RESID 238:256 )E238 - 256
27X-RAY DIFFRACTION27( CHAIN E AND RESID 257:269 )E257 - 269
28X-RAY DIFFRACTION28( CHAIN E AND RESID 270:284 )E270 - 284
29X-RAY DIFFRACTION29( CHAIN E AND RESID 285:322 )E285 - 322
30X-RAY DIFFRACTION30( CHAIN E AND RESID 323:336 )E323 - 336
31X-RAY DIFFRACTION31( CHAIN E AND RESID 337:354 )E337 - 354
32X-RAY DIFFRACTION32( CHAIN F AND RESID 138:142 )F138 - 142
33X-RAY DIFFRACTION33( CHAIN F AND RESID 143:190 )F143 - 190
34X-RAY DIFFRACTION34( CHAIN F AND RESID 191:235 )F191 - 235
35X-RAY DIFFRACTION35( CHAIN G AND RESID 150:178 )G150 - 178
36X-RAY DIFFRACTION36( CHAIN G AND RESID 179:212 )G179 - 212
37X-RAY DIFFRACTION37( CHAIN G AND RESID 213:222 )G213 - 222
38X-RAY DIFFRACTION38( CHAIN G AND RESID 223:269 )G223 - 269
39X-RAY DIFFRACTION39( CHAIN G AND RESID 270:304 )G270 - 304
40X-RAY DIFFRACTION40( CHAIN G AND RESID 305:322 )G305 - 322
41X-RAY DIFFRACTION41( CHAIN G AND RESID 323:353 )G323 - 353
42X-RAY DIFFRACTION42( CHAIN H AND RESID 140:178 )H140 - 178
43X-RAY DIFFRACTION43( CHAIN H AND RESID 179:202 )H179 - 202
44X-RAY DIFFRACTION44( CHAIN H AND RESID 203:235 )H203 - 235
45X-RAY DIFFRACTION45( CHAIN I AND RESID 151:163 )I151 - 163
46X-RAY DIFFRACTION46( CHAIN I AND RESID 164:188 )I164 - 188
47X-RAY DIFFRACTION47( CHAIN I AND RESID 189:202 )I189 - 202
48X-RAY DIFFRACTION48( CHAIN I AND RESID 203:237 )I203 - 237
49X-RAY DIFFRACTION49( CHAIN I AND RESID 238:269 )I238 - 269
50X-RAY DIFFRACTION50( CHAIN I AND RESID 270:295 )I270 - 295
51X-RAY DIFFRACTION51( CHAIN I AND RESID 296:314 )I296 - 314
52X-RAY DIFFRACTION52( CHAIN I AND RESID 315:353 )I315 - 353
53X-RAY DIFFRACTION53( CHAIN J AND RESID 138:142 )J138 - 142
54X-RAY DIFFRACTION54( CHAIN J AND RESID 143:178 )J143 - 178
55X-RAY DIFFRACTION55( CHAIN J AND RESID 179:235 )J179 - 235
56X-RAY DIFFRACTION56( CHAIN K AND RESID 150:202 )K150 - 202
57X-RAY DIFFRACTION57( CHAIN K AND RESID 203:245 )K203 - 245
58X-RAY DIFFRACTION58( CHAIN K AND RESID 246:304 )K246 - 304
59X-RAY DIFFRACTION59( CHAIN K AND RESID 305:325 )K305 - 325
60X-RAY DIFFRACTION60( CHAIN K AND RESID 326:354 )K326 - 354
61X-RAY DIFFRACTION61( CHAIN L AND RESID 140:170 )L140 - 170
62X-RAY DIFFRACTION62( CHAIN L AND RESID 171:178 )L171 - 178
63X-RAY DIFFRACTION63( CHAIN L AND RESID 179:189 )L179 - 189
64X-RAY DIFFRACTION64( CHAIN L AND RESID 190:207 )L190 - 207
65X-RAY DIFFRACTION65( CHAIN L AND RESID 208:224 )L208 - 224
66X-RAY DIFFRACTION66( CHAIN L AND RESID 225:235 )L225 - 235
67X-RAY DIFFRACTION67( CHAIN M AND RESID 151:269 )M151 - 269
68X-RAY DIFFRACTION68( CHAIN M AND RESID 270:354 )M270 - 354
69X-RAY DIFFRACTION69( CHAIN N AND RESID 141:178 )N141 - 178
70X-RAY DIFFRACTION70( CHAIN N AND RESID 179:202 )N179 - 202
71X-RAY DIFFRACTION71( CHAIN N AND RESID 203:235 )N203 - 235
72X-RAY DIFFRACTION72( CHAIN O AND RESID 150:176 )O150 - 176
73X-RAY DIFFRACTION73( CHAIN O AND RESID 177:188 )O177 - 188
74X-RAY DIFFRACTION74( CHAIN O AND RESID 189:202 )O189 - 202
75X-RAY DIFFRACTION75( CHAIN O AND RESID 203:222 )O203 - 222
76X-RAY DIFFRACTION76( CHAIN O AND RESID 223:256 )O223 - 256
77X-RAY DIFFRACTION77( CHAIN O AND RESID 257:269 )O257 - 269
78X-RAY DIFFRACTION78( CHAIN O AND RESID 270:295 )O270 - 295
79X-RAY DIFFRACTION79( CHAIN O AND RESID 296:314 )O296 - 314
80X-RAY DIFFRACTION80( CHAIN O AND RESID 315:338 )O315 - 338
81X-RAY DIFFRACTION81( CHAIN O AND RESID 339:354 )O339 - 354
82X-RAY DIFFRACTION82( CHAIN P AND RESID 141:162 )P141 - 162
83X-RAY DIFFRACTION83( CHAIN P AND RESID 163:178 )P163 - 178
84X-RAY DIFFRACTION84( CHAIN P AND RESID 179:190 )P179 - 190
85X-RAY DIFFRACTION85( CHAIN P AND RESID 191:202 )P191 - 202
86X-RAY DIFFRACTION86( CHAIN P AND RESID 203:235 )P203 - 235
87X-RAY DIFFRACTION87( CHAIN Q AND RESID 151:230 )Q151 - 230
88X-RAY DIFFRACTION88( CHAIN Q AND RESID 231:295 )Q231 - 295
89X-RAY DIFFRACTION89( CHAIN Q AND RESID 296:314 )Q296 - 314
90X-RAY DIFFRACTION90( CHAIN Q AND RESID 315:354 )Q315 - 354
91X-RAY DIFFRACTION91( CHAIN R AND RESID 141:170 )R141 - 170
92X-RAY DIFFRACTION92( CHAIN R AND RESID 171:190 )R171 - 190
93X-RAY DIFFRACTION93( CHAIN R AND RESID 191:202 )R191 - 202
94X-RAY DIFFRACTION94( CHAIN R AND RESID 203:235 )R203 - 235
95X-RAY DIFFRACTION95( CHAIN S AND RESID 151:178 )S151 - 178
96X-RAY DIFFRACTION96( CHAIN S AND RESID 179:202 )S179 - 202
97X-RAY DIFFRACTION97( CHAIN S AND RESID 203:237 )S203 - 237
98X-RAY DIFFRACTION98( CHAIN S AND RESID 238:256 )S238 - 256
99X-RAY DIFFRACTION99( CHAIN S AND RESID 257:269 )S257 - 269
100X-RAY DIFFRACTION100( CHAIN S AND RESID 270:295 )S270 - 295
101X-RAY DIFFRACTION101( CHAIN S AND RESID 296:311 )S296 - 311
102X-RAY DIFFRACTION102( CHAIN S AND RESID 312:321 )S312 - 321
103X-RAY DIFFRACTION103( CHAIN S AND RESID 322:336 )S322 - 336
104X-RAY DIFFRACTION104( CHAIN S AND RESID 337:353 )S337 - 353
105X-RAY DIFFRACTION105( CHAIN T AND RESID 141:178 )T141 - 178
106X-RAY DIFFRACTION106( CHAIN T AND RESID 179:190 )T179 - 190
107X-RAY DIFFRACTION107( CHAIN T AND RESID 191:202 )T191 - 202
108X-RAY DIFFRACTION108( CHAIN T AND RESID 203:235 )T203 - 235

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