PDB-3m6s: Crystal structure of H1N1pdm Hemagglutinin

ID or keywords:

Entry

Database: PDB / ID: 3m6s

Title

Crystal structure of H1N1pdm Hemagglutinin

Components

(Hemagglutinin

) x 2

Keywords

VIRAL PROTEIN /

Influenza virus /

Hemagglutinin /

pandemic /

Envelope protein /

Fusion protein / Host cell membrane / Host membrane /

Membrane /

Transmembrane /

Virion

Function / homology

Function and homology information

clathrin-dependent endocytosis of virus by host cell / membrane => GO:0016020 / host cell surface receptor binding / apical plasma membrane / fusion of virus membrane with host plasma membrane / fusion of virus membrane with host endosome membrane /

viral envelope / virion attachment to host cell / host cell plasma membrane
Similarity search - Function

Biological species

Influenza A virus

Method

X-RAY DIFFRACTION /

SYNCHROTRON /

MOLECULAR REPLACEMENT / Resolution: 2.8 Å

Authors

Yang, H. / Carney, P.J. / Stevens, J.

Citation

Journal: PLoS Curr / Year: 2010
Title: Structure and Receptor binding properties of a pandemic H1N1 virus hemagglutinin
Authors: Yang, H. / Carney, P. / Stevens, J.

History

Deposition	Mar 16, 2010	Deposition site: RCSB / Processing site: RCSB
Revision 1.0	Mar 31, 2010	Provider: repository / Type: Initial release
Revision 1.1	Jul 13, 2011	Group: Advisory / Refinement description / Version format compliance
Revision 1.2	Mar 19, 2014	Group: Database references
Revision 2.0	Jul 29, 2020	Group: Advisory / Atomic model ...Advisory / Atomic model / Data collection / Database references / Derived calculations / Structure summary Category: atom_site / chem_comp ...atom_site / chem_comp / database_PDB_caveat / entity / pdbx_branch_scheme / pdbx_chem_comp_identifier / pdbx_entity_branch / pdbx_entity_branch_descriptor / pdbx_entity_branch_link / pdbx_entity_branch_list / pdbx_entity_nonpoly / pdbx_nonpoly_scheme / pdbx_struct_assembly_gen / pdbx_unobs_or_zero_occ_atoms / pdbx_validate_chiral / pdbx_validate_close_contact / struct_asym / struct_conn / struct_ref_seq_dif / struct_site / struct_site_gen Item: _atom_site.B_iso_or_equiv / _atom_site.Cartn_x ..._atom_site.B_iso_or_equiv / _atom_site.Cartn_x / _atom_site.Cartn_y / _atom_site.Cartn_z / _atom_site.auth_asym_id / _atom_site.auth_atom_id / _atom_site.auth_comp_id / _atom_site.auth_seq_id / _atom_site.label_asym_id / _atom_site.label_atom_id / _atom_site.label_comp_id / _atom_site.label_entity_id / _atom_site.type_symbol / _chem_comp.name / _chem_comp.type / _pdbx_struct_assembly_gen.asym_id_list / _pdbx_validate_chiral.auth_asym_id / _pdbx_validate_chiral.auth_seq_id / _pdbx_validate_close_contact.auth_asym_id_2 / _pdbx_validate_close_contact.auth_seq_id_2 / _struct_conn.pdbx_dist_value / _struct_conn.pdbx_leaving_atom_flag / _struct_conn.pdbx_role / _struct_conn.ptnr1_auth_asym_id / _struct_conn.ptnr1_auth_comp_id / _struct_conn.ptnr1_auth_seq_id / _struct_conn.ptnr1_label_asym_id / _struct_conn.ptnr1_label_atom_id / _struct_conn.ptnr1_label_comp_id / _struct_conn.ptnr1_label_seq_id / _struct_conn.ptnr2_auth_asym_id / _struct_conn.ptnr2_auth_comp_id / _struct_conn.ptnr2_auth_seq_id / _struct_conn.ptnr2_label_asym_id / _struct_conn.ptnr2_label_comp_id / _struct_ref_seq_dif.details Description: Carbohydrate remediation / Provider: repository / Type: Remediation
Revision 2.1	Oct 6, 2021	Group: Advisory / Database references / Structure summary Category: chem_comp / database_2 ...chem_comp / database_2 / pdbx_unobs_or_zero_occ_atoms / struct_ref_seq_dif Item: _chem_comp.pdbx_synonyms / _database_2.pdbx_DOI ..._chem_comp.pdbx_synonyms / _database_2.pdbx_DOI / _database_2.pdbx_database_accession / _struct_ref_seq_dif.details
Revision 2.2	Sep 6, 2023	Group: Data collection / Refinement description Category: chem_comp_atom / chem_comp_bond / pdbx_initial_refinement_model

Structure viewer	Molecule: MolmilJmol/JSmol

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Validation report

Arichive directory	https://data.pdbj.org/pub/pdb/validation_reports/m6/3m6s ftp://data.pdbj.org/pub/pdb/validation_reports/m6/3m6s	HTTPS FTP

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Related structure data

Related structure data	3ku3S S: Starting model for refinement
Similar structure data	3ubn-2 3ubj-2 5bqz-d 2fk0-3 4lxv-2 4yya-1 4wss-1 3ubq-1 5bqy-d 2wr4-d 4jtv-1 5t08-d 10696 5bny-d 3qqo-d 3ubj-1 3ube-2 2wr2-d 3ubn-1 6v47-d Search similar-shape structures by Omokage search (details) Search similar-shape structures by Omokage search (details) Search similar-shape structures by Omokage search (details) Similarity search - Function & homologyF&H Search Similarity search - FunctionF&H Search Similarity search - HomologyF&H Search

Related structure data

3ku3S

S: Starting model for refinement

Similar structure data

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Links

PDB pages	PDBj / wwPDB / NCBI
Related items in Molecule of the Month	#76 - Apr 2006 Hemagglutinin similarity (6) #170 - Feb 2014 Broadly Neutralizing Antibodies similarity (5) #259 - Jul 2021 Designed Proteins and Citizen Science similarity (5) #62 - Feb 2005 Major Histocompatibility Complex similarity (5) #63 - Mar 2005 T-Cell Receptor similarity (5) #200 - Aug 2016 Quasisymmetry in Icosahedral Viruses similarity (1)

Deposited unit

A: Hemagglutinin

B: Hemagglutinin

C: Hemagglutinin

D: Hemagglutinin

E: Hemagglutinin

F: Hemagglutinin

G: Hemagglutinin

H: Hemagglutinin

I: Hemagglutinin

J: Hemagglutinin

K: Hemagglutinin

L: Hemagglutinin

hetero molecules

348 kDa, 12 polymers
Search similar-shape structures of this assembly by Omokage search (details)

1

A: Hemagglutinin

B: Hemagglutinin

C: Hemagglutinin

D: Hemagglutinin

E: Hemagglutinin

F: Hemagglutinin

hetero molecules

defined by author&software
173 kDa, 6 polymers
Search similar-shape structures of this assembly by Omokage search (details)

2

G: Hemagglutinin

H: Hemagglutinin

hetero molecules

I: Hemagglutinin

J: Hemagglutinin

K: Hemagglutinin

L: Hemagglutinin

hetero molecules

defined by author&software
175 kDa, 6 polymers
Search similar-shape structures of this assembly by Omokage search (details)

Unit cell

Length a, b, c (Å)	73.984, 109.709, 129.896
Angle α, β, γ (deg.)	86.25, 74.68, 75.10
Int Tables number	1
Space group name H-M	P1

Noncrystallographic symmetry (NCS)

NCS domain:

ID	Ens-ID	Details
1	1	F
2	1	D
3	1	B
4	1	H
5	1	J
6	1	L
1	2	E
2	2	C
3	2	G
4	2	A
5	2	I
6	2	K

NCS ensembles :

ID
1
2

Details

The biological unit is a trimer.

#1: Protein

Hemagglutinin /

Mass: 36661.324 Da / Num. of mol.: 6 / Fragment: Hemagglutinin HA1
Source method: isolated from a genetically manipulated source
Source: (gene. exp.)

Influenza A virus / Strain: A/Darwin/2001/2009(H1N1) / Gene: HA / Production host:

Trichoplusia ni (cabbage looper) / References: UniProt: C5MV42

#2: Protein

Hemagglutinin /

Mass: 20759.008 Da / Num. of mol.: 6 / Fragment: Hemagglutinin HA2
Source method: isolated from a genetically manipulated source
Source: (gene. exp.)

Influenza A virus / Strain: A/Darwin/2001/2009(H1N1) / Gene: HA / Production host:

Trichoplusia ni (cabbage looper) / References: UniProt: C5MV42

#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

Descriptor	Type	Program
DGlcpNAcb1-4DGlcpNAcb1-	Glycam Condensed Sequence	GMML 1.0
WURCS=2.0/1,2,1/[a2122h-1b_1-5_2*NCC/3=O]/1-1/a4-b1	WURCS	PDB2Glycan 1.1.0
[]{[(4+1)][b-D-GlcpNAc]{[(4+1)][b-D-GlcpNAc]{}}}	LINUCS	PDB-CARE

#4: Polysaccharide

alpha-D-mannopyranose-(1-4)-2-acetamido-2-deoxy-beta-D-glucopyranose-(1-4)-2-acetamido-2-deoxy-beta- ...

Type: oligosaccharide

/ Mass: 586.542 Da / Num. of mol.: 1
Source method: isolated from a genetically manipulated source

Descriptor	Type	Program
DManpa1-4DGlcpNAcb1-4DGlcpNAcb1-	Glycam Condensed Sequence	GMML 1.0
WURCS=2.0/2,3,2/[a2122h-1b_1-5_2*NCC/3=O][a1122h-1a_1-5]/1-1-2/a4-b1_b4-c1	WURCS	PDB2Glycan 1.1.0
[]{[(4+1)][b-D-GlcpNAc]{[(4+1)][b-D-GlcpNAc]{[(4+1)][b-D-Manp]{}}}}	LINUCS	PDB-CARE

#5: 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.: 4
Source method: isolated from a genetically manipulated source
Formula: C₈H₁₅NO₆

Identifier	Type	Program
DGlcpNAcb	CONDENSED IUPAC CARBOHYDRATE SYMBOL	GMML 1.0
N-acetyl-b-D-glucopyranosamine	COMMON NAME	GMML 1.0
b-D-GlcpNAc	IUPAC CARBOHYDRATE SYMBOL	PDB-CARE 1.0
GlcNAc	SNFG CARBOHYDRATE SYMBOL	GMML 1.0

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Experiment

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

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Sample preparation

Crystal	Density Matthews: 2.85 Å³/Da / Density % sol: 56.88 %
Crystal grow	Temperature: 293 K / Method: microbatch under oil / pH: 7.5 Details: 22% PEG2000MME, 0.1M HEPES at pH7.5, microbatch under oil, temperature 293K

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Data collection

Diffraction	Mean temperature: 100 K
Diffraction source	Source: SYNCHROTRON / Site: APS / Beamline: 22-BM / Wavelength: 1 Å
Detector	Type: MARMOSAIC 225 mm CCD / Detector: CCD / Date: Feb 10, 2010
Radiation	Protocol: SINGLE WAVELENGTH / Monochromatic (M) / Laue (L): M / Scattering type: x-ray
Radiation wavelength	Wavelength: 1 Å / Relative weight: 1
Reflection	Resolution: 2.8→50 Å / Num. all: 96521 / Num. obs: 94645 / % possible obs: 98.1 % / Redundancy: 2 % / R_sym value: 0.076
Reflection shell	Resolution: 2.8→2.9 Å / Redundancy: 1.9 % / Mean I/σ(I) obs: 1.6 / R_sym value: 0.442 / % possible all: 95.7

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Processing

Software

Name	Version	Classification
HKL-2000		data collection
PHASER		phasing
REFMAC	5.5.0102	refinement
HKL-2000		data reduction
HKL-2000		data scaling

Refinement

Method to determine structure:

MOLECULAR REPLACEMENT
Starting model: PDB entry 3KU3

3ku3

Resolution: 2.8→50 Å / Cor.coef. F_o:F_c: 0.912 / Cor.coef. F_o:F_c free: 0.889 / SU B: 36.705 / SU ML: 0.32 / TLS residual ADP flag: LIKELY RESIDUAL / Cross valid method: THROUGHOUT / ESU R Free: 0.381 / Stereochemistry target values: MAXIMUM LIKELIHOOD / Details: HYDROGENS HAVE BEEN ADDED IN THE RIDING POSITIONS

	R_factor	Num. reflection	% reflection	Selection details
R_free	0.25588	4608	5 %	RANDOM
R_work	0.22978	-	-	-
obs	0.23109	87344	98.09 %	-

Solvent computation

Ion probe radii: 0.8 Å / Shrinkage radii: 0.8 Å / VDW probe radii: 1.4 Å / Solvent model: MASK

Displacement parameters

B_iso mean: 24.604 Å²

	B_aniso -1	B_aniso -2	B_aniso -3
1-	-0.04 Å²	0.94 Å²	1.83 Å²
2-	-	2.55 Å²	-1.87 Å²
3-	-	-	-3.72 Å²

Refinement step

Cycle: LAST / Resolution: 2.8→50 Å

	Protein	Nucleic acid	Ligand	Solvent	Total
Num. atoms	23380	0	207	0	23587

Refine LS restraints

Refine-ID	Type	Dev ideal	Dev ideal target	Number
X-RAY DIFFRACTION	r_bond_refined_d	0.016	0.022	24181
X-RAY DIFFRACTION	r_bond_other_d
X-RAY DIFFRACTION	r_angle_refined_deg	1.701	1.951	32795
X-RAY DIFFRACTION	r_angle_other_deg
X-RAY DIFFRACTION	r_dihedral_angle_1_deg	7.077	5	2946
X-RAY DIFFRACTION	r_dihedral_angle_2_deg	36.776	25.062	1130
X-RAY DIFFRACTION	r_dihedral_angle_3_deg	20.275	15	4086
X-RAY DIFFRACTION	r_dihedral_angle_4_deg	21.462	15	84
X-RAY DIFFRACTION	r_chiral_restr	0.125	0.2	3565
X-RAY DIFFRACTION	r_gen_planes_refined	0.006	0.021	18340
X-RAY DIFFRACTION	r_gen_planes_other
X-RAY DIFFRACTION	r_nbd_refined
X-RAY DIFFRACTION	r_nbd_other
X-RAY DIFFRACTION	r_nbtor_refined
X-RAY DIFFRACTION	r_nbtor_other
X-RAY DIFFRACTION	r_xyhbond_nbd_refined
X-RAY DIFFRACTION	r_xyhbond_nbd_other
X-RAY DIFFRACTION	r_metal_ion_refined
X-RAY DIFFRACTION	r_metal_ion_other
X-RAY DIFFRACTION	r_symmetry_vdw_refined
X-RAY DIFFRACTION	r_symmetry_vdw_other
X-RAY DIFFRACTION	r_symmetry_hbond_refined
X-RAY DIFFRACTION	r_symmetry_hbond_other
X-RAY DIFFRACTION	r_symmetry_metal_ion_refined
X-RAY DIFFRACTION	r_symmetry_metal_ion_other
X-RAY DIFFRACTION	r_mcbond_it	0.66	1.5	14671
X-RAY DIFFRACTION	r_mcbond_other
X-RAY DIFFRACTION	r_mcangle_it	1.332	2	23685
X-RAY DIFFRACTION	r_scbond_it	2.107	3	9510
X-RAY DIFFRACTION	r_scangle_it	3.651	4.5	9110
X-RAY DIFFRACTION	r_rigid_bond_restr
X-RAY DIFFRACTION	r_sphericity_free
X-RAY DIFFRACTION	r_sphericity_bonded

Refine LS restraints NCS

Refine-ID: X-RAY DIFFRACTION

Ens-ID	Dom-ID	Auth asym-ID	Number	Type	Rms dev position (Å)	Weight position
1	1	F	676	tight positional	0.05	0.05
1	2	D	676	tight positional	0.06	0.05
1	3	B	676	tight positional	0.06	0.05
1	4	H	676	tight positional	0.06	0.05
1	5	J	676	tight positional	0.1	0.05
1	6	L	676	tight positional	0.06	0.05
2	1	E	1280	tight positional	0.05	0.05
2	2	C	1280	tight positional	0.06	0.05
2	3	G	1280	tight positional	0.07	0.05
2	4	A	1280	tight positional	0.07	0.05
2	5	I	1280	tight positional	0.06	0.05
2	6	K	1280	tight positional	0.07	0.05
1	1	F	687	loose positional	0.08	5
1	2	D	687	loose positional	0.08	5
1	3	B	687	loose positional	0.08	5
1	4	H	687	loose positional	0.07	5
1	5	J	687	loose positional	0.11	5
1	6	L	687	loose positional	0.07	5
2	1	E	1219	loose positional	0.06	5
2	2	C	1219	loose positional	0.07	5
2	3	G	1219	loose positional	0.08	5
2	4	A	1219	loose positional	0.08	5
2	5	I	1219	loose positional	0.06	5
2	6	K	1219	loose positional	0.08	5
1	1	F	676	tight thermal	0.08	0.5
1	2	D	676	tight thermal	0.09	0.5
1	3	B	676	tight thermal	0.09	0.5
1	4	H	676	tight thermal	0.09	0.5
1	5	J	676	tight thermal	0.1	0.5
1	6	L	676	tight thermal	0.09	0.5
2	1	E	1280	tight thermal	0.09	0.5
2	2	C	1280	tight thermal	0.11	0.5
2	3	G	1280	tight thermal	0.12	0.5
2	4	A	1280	tight thermal	0.1	0.5
2	5	I	1280	tight thermal	0.09	0.5
2	6	K	1280	tight thermal	0.1	0.5
1	1	F	687	loose thermal	0.09	10
1	2	D	687	loose thermal	0.11	10
1	3	B	687	loose thermal	0.09	10
1	4	H	687	loose thermal	0.09	10
1	5	J	687	loose thermal	0.13	10
1	6	L	687	loose thermal	0.1	10
2	1	E	1219	loose thermal	0.11	10
2	2	C	1219	loose thermal	0.13	10
2	3	G	1219	loose thermal	0.14	10
2	4	A	1219	loose thermal	0.11	10
2	5	I	1219	loose thermal	0.1	10
2	6	K	1219	loose thermal	0.12	10

LS refinement shell

Resolution: 2.8→2.873 Å / Total num. of bins used: 20

	R_factor	Num. reflection	% reflection
R_free	0.341	301	-
R_work	0.314	6322	-
obs	-	-	95.78 %

Refinement TLS params.

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

ID	L11 (°²)	L12 (°²)	L13 (°²)	L22 (°²)	L23 (°²)	L33 (°²)	S11 (Å °)	S12 (Å °)	S13 (Å °)	S21 (Å °)	S22 (Å °)	S23 (Å °)	S31 (Å °)	S32 (Å °)	S33 (Å °)	T11 (Å²)	T12 (Å²)	T13 (Å²)	T22 (Å²)	T23 (Å²)	T33 (Å²)	Origin x (Å)	Origin y (Å)	Origin z (Å)
1	0.2636	-0.1568	-0.091	0.3746	0.1917	3.1546	0.0699	0.0747	0.0871	-0.0013	0.018	-0.1042	-0.5865	0.385	-0.0878	0.2713	-0.1449	0.0706	0.163	-0.0505	0.2958	-20.3997	-12.5839	-34.3041
2	0.9741	0.3977	1.3252	1.4423	3.511	10.8841	-0.0265	-0.14	-0.0565	0.205	0.0674	-0.0523	-0.2095	-0.0649	-0.0409	0.5231	0.0516	0.0635	0.0441	0.0017	0.1584	-28.0342	-4.9711	17.3059
3	0.386	-0.0362	0.381	0.7952	1.2226	2.677	0.1411	-0.0291	-0.1563	0.0388	0.0894	0.054	0.0931	0.1175	-0.2306	0.1091	-0.0074	-0.032	0.0197	-0.0257	0.341	-36.6417	-42.1848	-28.5966
4	0.6682	0.0418	-0.3632	0.2329	0.4813	11.1244	0.181	-0.1284	-0.0831	0.0213	0.0328	0.0482	-0.5841	-0.7118	-0.2138	0.153	0.0332	0.0705	0.1154	0.0621	0.1983	-37.3934	-23.439	21.6924
5	0.5635	-0.1701	0.9074	0.409	0.0613	3.5393	0.0073	-0.1756	0.0162	-0.0016	0.1137	0.1646	-0.4748	-0.7234	-0.121	0.2363	0.1418	0.0438	0.2069	0.0829	0.3478	-54.4456	-13.7762	-35.4145
6	0.5362	-0.252	-1.1285	1.3503	1.9589	8.7406	0.1142	-0.2415	0.0403	0.1494	0.1865	0.0417	-1.3844	-0.6824	-0.3007	0.6315	0.3949	0.1445	0.4988	0.018	0.2205	-49.6245	-6.0546	17.599
7	0.9272	0.1156	0.8115	0.5906	0.0337	4.2825	0.0388	-0.1212	-0.1262	-0.0214	-0.0493	0.0762	0.4621	-0.1389	0.0106	0.0942	-0.018	-0.02	0.0242	-0.0058	0.191	-18.2701	-78.8485	-50.1068
8	1.4507	-0.1511	-1.8666	0.7984	-0.1121	7.8911	0.1193	-0.7443	0.0715	0.331	-0.0025	0.0096	-0.6766	0.8828	-0.1168	0.249	-0.102	-0.0338	0.523	-0.0295	0.1356	-10.8808	-69.6384	2.3853
9	0.7734	0.0489	0.2854	0.6542	0.5977	4.3455	0.0507	0.1578	-0.262	-0.0912	-0.0045	-0.2932	0.521	1.3717	-0.0461	0.1727	0.2353	0.0523	0.6376	0.0248	0.3189	-58.0768	-80.9825	-50.4158
10	0.4003	-0.0977	2.3569	0.2171	-0.6858	15.2046	-0.0458	0.0747	0.0219	0.0253	-0.1504	-0.1252	-0.0006	0.2972	0.1962	0.0749	-0.0176	-0.0459	0.5596	0.0756	0.1171	-63.3914	-69.8712	2.2438
11	0.7897	0.145	0.2544	0.856	0.6237	5.2118	-0.0768	0.0577	0.2107	-0.1864	0.1412	-0.035	-0.6252	0.6515	-0.0644	0.1408	-0.1589	-0.0217	0.1959	0.0008	0.2465	-73.4443	-50.5896	-53.8326
12	1.668	0.3501	0.4514	1.6197	1.6739	6.5169	0.1842	-0.8433	0.25	0.7925	-0.1554	-0.0653	0.3796	0.5687	-0.0289	0.4566	-0.1862	-0.0701	0.6072	-0.1204	0.1868	-73.3476	-51.6933	0.2944

Refinement TLS group

ID	Refine-ID	Refine TLS-ID	Auth asym-ID	Auth seq-ID
1	X-RAY DIFFRACTION	1	A	-2 - 322
2	X-RAY DIFFRACTION	1	A	328
3	X-RAY DIFFRACTION	2	B	2 - 170
4	X-RAY DIFFRACTION	3	C	0 - 322
5	X-RAY DIFFRACTION	3	C	328 - 329
6	X-RAY DIFFRACTION	4	D	2 - 172
7	X-RAY DIFFRACTION	5	E	0 - 322
8	X-RAY DIFFRACTION	5	E	328
9	X-RAY DIFFRACTION	6	F	2 - 172
10	X-RAY DIFFRACTION	7	G	1 - 322
11	X-RAY DIFFRACTION	7	G	328 - 332
12	X-RAY DIFFRACTION	8	H	2 - 171
13	X-RAY DIFFRACTION	9	I	0 - 321
14	X-RAY DIFFRACTION	9	I	328 - 329
15	X-RAY DIFFRACTION	10	J	2 - 171
16	X-RAY DIFFRACTION	11	K	2 - 322
17	X-RAY DIFFRACTION	11	K	328 - 331
18	X-RAY DIFFRACTION	12	L	2 - 172

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