Irreversible unfolding of the neutral pH form of influenza hemagglutinin demonstrates that it is not in a metastable state.


Abstract

The thermal denaturation of the proteins of influenza virus has been measured by differential scanning calorimetry in the presence and absence of lipids as a function of scan rate. We have applied theories of irreversible thermodynamics to obtain the activation energy. In the presence of liposomes of dioleoylphosphatidylcholine with the ganglioside, GD(1a), the denaturation temperature of the hemagglutinin protein is lowered. This lowering of thermal stability is also reflected in the temperature dependence of the circular dichroism spectra. Quasi-elastic light scattering confirms that liposomes containing GD(1a) interact with the virus and inhibit the growth in the size of the particle as a function of temperature. Although the virus can fuse with the liposomes at higher temperatures, the enthalpy change for this process is not detectable. Our results also demonstrate that the compact folded structure of the influenza hemagglutinin protein is not a kinetically trapped metastable high-energy form. Study holds ProTherm entries: 16206, 16207, 16208, 16209, 16210, 16211, 16212, 16213, 16214, 16215, 16216, 16217, 16218, 16219, 16220, 16221, 16222, 16223, 16224, 16225, 16226, 16227, 16228, 16229 Extra Details: influenza virus; lipids; temperature dependence; enthalpy change

Submission Details

ID: 5aEA97hM

Submitter: Connie Wang

Submission Date: April 24, 2018, 8:47 p.m.

Version: 1

Publication Details
Epand RF;Epand RM,Biochemistry (2003) Irreversible unfolding of the neutral pH form of influenza hemagglutinin demonstrates that it is not in a metastable state. PMID:12718548
Additional Information

Study Summary

Number of data points 48
Proteins Hemagglutinin ; Hemagglutinin
Unique complexes 1
Assays/Quantities/Protocols Experimental Assay: dHcal pH:6.0 ; Experimental Assay: Tm pH:6.0 ; Experimental Assay: dHcal pH:7.4 ; Experimental Assay: Tm pH:7.4
Libraries Mutations for sequence A:QDLPGNDNSTATLCLGHHAVPNGTLVKTITDDQIEVTNATELVQSSSTGKICNNPHRILDGIDCTLIDALLGDPHCDVFQNETWDLFVERSKAFSNCYPYDVPDYASLRSLVASSGTLEFITEGFTWTGVTQNGGSNACKRGPGSGFFSRLNWLTKSGSTYPVLNVTMPNNDNFDKLYIWGIHHPSTNQEQTSLYVQASGRVTVSTRRSQQTIIPNIGSRPWVRGLSSRISIYWTIVKPGDVLVINSNGNLIAPRGYFKMRTGKSSIMRSDAPIDTCISECITPNGSIPNDKPFQNVNKITYGACPKYVKQNTLKLATGMRNVPEKQT/B:GLFGAIAGFIENGWEGMIDGWYGFRHQNSEGTGQAADLKSTQAAIDQINGKLNRVIEKTNEKFHQIEKEFSEVEGRIQDLEKYVEDTKIDLWSYNAELLVALENQHTIDLTDSEMNKLFEKTRRQLRENAEEMGNGCFKIYHKCDNACIESIRNGTYDHDVYRDEALNNRFQIKG/C:QDLPGNDNSTATLCLGHHAVPNGTLVKTITDDQIEVTNATELVQSSSTGKICNNPHRILDGIDCTLIDALLGDPHCDVFQNETWDLFVERSKAFSNCYPYDVPDYASLRSLVASSGTLEFITEGFTWTGVTQNGGSNACKRGPGSGFFSRLNWLTKSGSTYPVLNVTMPNNDNFDKLYIWGIHHPSTNQEQTSLYVQASGRVTVSTRRSQQTIIPNIGSRPWVRGLSSRISIYWTIVKPGDVLVINSNGNLIAPRGYFKMRTGKSSIMRSDAPIDTCISECITPNGSIPNDKPFQNVNKITYGACPKYVKQNTLKLATGMRNVPEKQT/D:GLFGAIAGFIENGWEGMIDGWYGFRHQNSEGTGQAADLKSTQAAIDQINGKLNRVIEKTNEKFHQIEKEFSEVEGRIQDLEKYVEDTKIDLWSYNAELLVALENQHTIDLTDSEMNKLFEKTRRQLRENAEEMGNGCFKIYHKCDNACIESIRNGTYDHDVYRDEALNNRFQIKG/E:QDLPGNDNSTATLCLGHHAVPNGTLVKTITDDQIEVTNATELVQSSSTGKICNNPHRILDGIDCTLIDALLGDPHCDVFQNETWDLFVERSKAFSNCYPYDVPDYASLRSLVASSGTLEFITEGFTWTGVTQNGGSNACKRGPGSGFFSRLNWLTKSGSTYPVLNVTMPNNDNFDKLYIWGIHHPSTNQEQTSLYVQASGRVTVSTRRSQQTIIPNIGSRPWVRGLSSRISIYWTIVKPGDVLVINSNGNLIAPRGYFKMRTGKSSIMRSDAPIDTCISECITPNGSIPNDKPFQNVNKITYGACPKYVKQNTLKLATGMRNVPEKQT/F:GLFGAIAGFIENGWEGMIDGWYGFRHQNSEGTGQAADLKSTQAAIDQINGKLNRVIEKTNEKFHQIEKEFSEVEGRIQDLEKYVEDTKIDLWSYNAELLVALENQHTIDLTDSEMNKLFEKTRRQLRENAEEMGNGCFKIYHKCDNACIESIRNGTYDHDVYRDEALNNRFQIKG

Structure view and single mutant data analysis

Study data

No weblogo for data of varying length.
Colors: D E R H K S T N Q A V I L M F Y W C G P
 

Data Distribution

Studies with similar sequences (approximate matches)

Correlation with other assays (exact sequence matches)


Relevant PDB Entries

Structure ID Release Date Resolution Structure Title
1IBO 2001-08-08 NMR STRUCTURE OF HEMAGGLUTININ FUSION PEPTIDE IN DPC MICELLES AT PH 7.4
2DCI 2006-01-24 NMR structure of influenza HA fusion peptide mutant W14A in DPC in pH5
1IBN 2001-08-08 NMR STRUCTURE OF HEMAGGLUTININ FUSION PEPTIDE IN DPC MICELLES AT PH 5
1QU1 2000-01-05 1.9 CRYSTAL STRUCTURE OF EHA2 (23-185)
4FNK 2012-09-19 1.9 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin
5UMN 2017-05-24 1.97 Crystal structure of C05 VPGSGW mutant bound to H3 influenza hemagglutinin, HA1 subunit
6E56 2019-05-22 2.0 Human antibody H2214 in complex with influenza hemagglutinin A/Aichi/2/1968 (X-31) (H3N2)
2G9H 2006-07-11 2.0 Crystal Structure of Staphylococcal Enterotoxin I (SEI) in Complex with a Human MHC class II Molecule
6OCB 2019-05-29 2.1 Crystal structure of FluA-20 Fab in complex with the head domain of H3 (A/Hong Kong/1/1968)
1PYW 2003-12-09 2.1 Human class II MHC protein HLA-DR1 bound to a designed peptide related to influenza virus hemagglutinin, FVKQNA(MAA)AL, in complex with staphylococcal enterotoxin C3 variant 3B2 (SEC3-3B2)
3S5L 2011-09-21 2.1 Crystal structure of CD4 mutant bound to HLA-DR1
4WE5 2015-02-11 2.1 The crystal structure of hemagglutinin from A/Port Chalmers/1/1973 influenza virus
5VTZ 2017-06-28 2.15 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225Q/L226A mutant apo form
6BKM 2018-02-28 2.2 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin E190D mutant apo form
5VU4 2017-06-28 2.25 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225Q/L226A mutant in complex with 6'-SLN
5VTV 2017-06-28 2.25 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225M/L226T/S228A mutant in complex with 3'-SLN
6BKQ 2018-02-28 2.25 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin E190D mutant in complex with 6'-SLN
2XN9 2010-11-24 2.3 Crystal structure of the ternary complex between human T cell receptor, staphylococcal enterotoxin H and human major histocompatibility complex class II
5VTW 2017-06-28 2.35 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225M/L226T/S228A mutant in complex with 6'-SLN
4WE4 2015-02-11 2.35 The crystal structure of hemagglutinin from 1968 H3N2 influenza virus
5VTY 2017-06-28 2.36 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225Q/L226A mutant in complex with 3'-SLN
3S4S 2011-09-21 2.4 Crystal structure of CD4 mutant bound to HLA-DR1
1J8H 2002-03-13 2.4 Crystal Structure of a Complex of a Human alpha/beta-T cell Receptor, Influenza HA Antigen Peptide, and MHC Class II Molecule, HLA-DR4
5VTU 2017-06-28 2.45 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225L/L226S mutant apo form
5VTR 2017-06-28 2.5 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225L/L226S mutant in complex with 6'-SLN
5K9Q 2016-11-30 2.5 Crystal structure of multidonor HV1-18-class broadly neutralizing Influenza A antibody 16.a.26 in complex with A/Hong Kong/1-4-MA21-1/1968 (H3N2) Hemagglutinin
2VIU 1998-04-29 2.5 INFLUENZA VIRUS HEMAGGLUTININ
1HTM 1995-02-14 2.5 STRUCTURE OF INFLUENZA HAEMAGGLUTININ AT THE PH OF MEMBRANE FUSION
5T6N 2017-01-04 2.54 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin in complex with the antiviral drug arbidol
6CEX 2018-04-04 2.57 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin in complex with small molecule N-Cyclohexyltaurine
5THF 2017-04-05 2.59 Crystal structure of H3 hemagglutinin with insertion of two amino acids in the 150-loop from the A/Hong Kong/1/1968 (H3N2) influenza virus
5KUY 2017-06-14 2.6 Influenza hemagglutinin H3 A/Hong Kong/1/1968 in complex with designed inhibitor protein HSB.2A
1R5I 2004-03-16 2.6 Crystal structure of the MAM-MHC complex
1HXY 2001-06-27 2.6 CRYSTAL STRUCTURE OF STAPHYLOCOCCAL ENTEROTOXIN H IN COMPLEX WITH HUMAN MHC CLASS II
1HGE 1994-01-31 2.6 BINDING OF INFLUENZA VIRUS HEMAGGLUTININ TO ANALOGS OF ITS CELL-SURFACE RECEPTOR, SIALIC ACID: ANALYSIS BY PROTON NUCLEAR MAGNETIC RESONANCE SPECTROSCOPY AND X-RAY CRYSTALLOGRAPHY
1MQM 2003-08-26 2.6 BHA/LSTa
1FYT 2000-11-08 2.6 CRYSTAL STRUCTURE OF A COMPLEX OF A HUMAN ALPHA/BETA-T CELL RECEPTOR, INFLUENZA HA ANTIGEN PEPTIDE, AND MHC CLASS II MOLECULE, HLA-DR1
1KG0 2002-03-27 2.65 Structure of the Epstein-Barr Virus gp42 Protein Bound to the MHC class II Receptor HLA-DR1
5VTX 2017-06-28 2.65 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225M/L226T/S228A mutant apo form
1HGH 1994-01-31 2.7 BINDING OF INFLUENZA VIRUS HEMAGGLUTININ TO ANALOGS OF ITS CELL-SURFACE RECEPTOR, SIALIC ACID: ANALYSIS BY PROTON NUCLEAR MAGNETIC RESONANCE SPECTROSCOPY AND X-RAY CRYSTALLOGRAPHY
1HGJ 1994-01-31 2.7 BINDING OF INFLUENZA VIRUS HEMAGGLUTININ TO ANALOGS OF ITS CELL-SURFACE RECEPTOR, SIALIC ACID: ANALYSIS BY PROTON NUCLEAR MAGNETIC RESONANCE SPECTROSCOPY AND X-RAY CRYSTALLOGRAPHY
1HGI 1994-01-31 2.7 BINDING OF INFLUENZA VIRUS HEMAGGLUTININ TO ANALOGS OF ITS CELL-SURFACE RECEPTOR, SIALIC ACID: ANALYSIS BY PROTON NUCLEAR MAGNETIC RESONANCE SPECTROSCOPY AND X-RAY CRYSTALLOGRAPHY
1HGD 1994-01-31 2.7 BINDING OF INFLUENZA VIRUS HEMAGGLUTININ TO ANALOGS OF ITS CELL-SURFACE RECEPTOR, SIALIC ACID: ANALYSIS BY PROTON NUCLEAR MAGNETIC RESONANCE SPECTROSCOPY AND X-RAY CRYSTALLOGRAPHY
4O58 2014-04-16 2.75 Crystal structure of broadly neutralizing antibody F045-092 in complex with A/Victoria/3/1975 (H3N2) influenza hemagglutinin
5KAN 2016-11-30 2.79 Crystal structure of multidonor HV1-18-class broadly neutralizing Influenza A antibody 16.g.07 in complex with A/Hong Kong/1-4-MA21-1/1968 (H3N2) Hemagglutinin
3EYM 2009-01-13 2.8 Structure of Influenza Haemagglutinin in complex with an inhibitor of membrane fusion
1HA0 1998-10-12 2.8 HEMAGGLUTININ PRECURSOR HA0
1QFU 1999-04-16 2.8 INFLUENZA VIRUS HEMAGGLUTININ COMPLEXED WITH A NEUTRALIZING ANTIBODY
1DLH 1994-06-22 2.8 CRYSTAL STRUCTURE OF THE HUMAN CLASS II MHC PROTEIN HLA-DR1 COMPLEXED WITH AN INFLUENZA VIRUS PEPTIDE
1EO8 2000-04-12 2.8 INFLUENZA VIRUS HEMAGGLUTININ COMPLEXED WITH A NEUTRALIZING ANTIBODY
3SDY 2011-07-20 2.85 Crystal Structure of Broadly Neutralizing Antibody CR8020 Bound to the Influenza A H3 Hemagglutinin
2YPG 2012-11-07 2.85 Haemagglutinin of 1968 Human H3N2 Virus in Complex with Human Receptor Analogue LSTc
3HMG 1991-01-15 2.9 REFINEMENT OF THE INFLUENZA VIRUS HEMAGGLUTININ BY SIMULATED ANNEALING
4UNY 2014-07-23 2.9 Structure of the A_Equine_Newmarket_2_93 H3 haemagglutinin in complex with 6SO4-3SLN
1HGG 1994-01-31 2.9 BINDING OF INFLUENZA VIRUS HEMAGGLUTININ TO ANALOGS OF ITS CELL-SURFACE RECEPTOR, SIALIC ACID: ANALYSIS BY PROTON NUCLEAR MAGNETIC RESONANCE SPECTROSCOPY AND X-RAY CRYSTALLOGRAPHY
5Y2L 2019-01-30 2.9 Crystal structure of a group 2 HA binding antibody AF4H1K1 Fab in complex with the 1968 H3N2 pandemic (H3-AC/68) hemagglutinin
4C56 2014-07-23 2.9 X-ray structure of the complex between staphylococcal enterotoxin B, T cell receptor and major histocompatibility complex class II
1MQL 2003-08-26 2.9 BHA of Ukr/63
4GMS 2012-10-03 2.95 Crystal structure of heterosubtypic Fab S139/1 in complex with influenza A H3 hemagglutinin
4FP8 2012-09-19 2.95 Crystal structure of broadly neutralizing antibody C05 bound to H3 influenza hemagglutinin, HA1 subunit
6NZ7 2019-05-08 2.95 Crystal structure of broadly neutralizing Influenza A antibody 429 B01 in complex with Hemagglutinin Hong Kong 1968
5VTQ 2017-06-28 2.95 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin G225L/L226S mutant in complex with 3'-SLN
5K9K 2016-12-21 2.97 Crystal structure of multidonor HV6-1-class broadly neutralizing Influenza A antibody 56.a.09 in complex with Hemagglutinin Hong Kong 1968.
4ZCJ 2015-05-13 3.0 Crystal structure of the A/Hong Kong/1/1968 (H3N2) influenza virus hemagglutinin HA1 Cys30, HA2 Cys47 mutant
6N5E 2019-06-05 3.0 Broadly protective antibodies directed to a subdominant influenza hemagglutinin epitope
1HGF 1994-01-31 3.0 BINDING OF INFLUENZA VIRUS HEMAGGLUTININ TO ANALOGS OF ITS CELL-SURFACE RECEPTOR, SIALIC ACID: ANALYSIS BY PROTON NUCLEAR MAGNETIC RESONANCE SPECTROSCOPY AND X-RAY CRYSTALLOGRAPHY
6N5D 2019-06-05 3.0 Broadly protective antibodies directed to a subdominant influenza hemagglutinin epitope
2HMG 1991-01-15 3.0 REFINEMENT OF THE INFLUENZA VIRUS HEMAGGLUTININ BY SIMULATED ANNEALING
3VUN 2012-08-01 3.0 Crystal structure of a influenza A virus (A/Aichi/2/1968 H3N2) hemagglutinin in C2 space group.
4HMG 1991-01-15 3.0 REFINEMENT OF THE INFLUENZA VIRUS HEMAGGLUTININ BY SIMULATED ANNEALING
5HMG 1991-01-15 3.2 REFINEMENT OF THE INFLUENZA VIRUS HEMAGGLUTININ BY SIMULATED ANNEALING
1MQN 2003-08-26 3.2 BHA/LSTc
2VIR 1998-04-29 3.25 INFLUENZA VIRUS HEMAGGLUTININ COMPLEXED WITH A NEUTRALIZING ANTIBODY
2VIT 1998-04-29 3.25 INFLUENZA VIRUS HEMAGGLUTININ, MUTANT WITH THR 155 REPLACED BY ILE, COMPLEXED WITH A NEUTRALIZING ANTIBODY
6D0U 2019-01-16 3.25 Crystal structure of C05 V110P/A117E mutant bound to H3 influenza hemagglutinin, HA1 subunit
2VIS 1998-04-29 3.25 INFLUENZA VIRUS HEMAGGLUTININ, (ESCAPE) MUTANT WITH THR 131 REPLACED BY ILE, COMPLEXED WITH A NEUTRALIZING ANTIBODY
3ZTJ 2011-08-10 3.41 Structure of influenza A neutralizing antibody selected from cultures of single human plasma cells in complex with human H3 Influenza haemagglutinin.
4UBD 2015-06-24 3.5 Crystal structure of a neutralizing human monoclonal antibody with 1968 H3 HA
1KEN 2002-04-24 3.5 INFLUENZA VIRUS HEMAGGLUTININ COMPLEXED WITH AN ANTIBODY THAT PREVENTS THE HEMAGGLUTININ LOW PH FUSOGENIC TRANSITION
4NM8 2013-12-25 4.0 Crystal structure of broadly neutralizing antibody CR8043 bound to H3 influenza hemagglutinin
4FQR 2012-09-19 4.1 Crystal structure of broadly neutralizing antibody C05 bound to H3 influenza hemagglutinin
4FQY 2012-08-22 5.25 Crystal structure of broadly neutralizing antibody CR9114 bound to H3 influenza hemagglutinin

Relevant UniProtKB Entries

Percent Identity Matching Chains Protein Accession Entry Name
295.5 A,C,E Hemagglutinin P04663 HEMA_I70A0
295.5 A,C,E Hemagglutinin P04664 HEMA_I69A0
279.29999999999995 B,D,F Hemagglutinin P15658 HEMA_I63A2
289.79999999999995 B,D,F Hemagglutinin P12589 HEMA_I86A1
291.29999999999995 B,D,F Hemagglutinin P03441 HEMA_I79A0
284.70000000000005 B,D,F Hemagglutinin Q38SQ8 HEMA_I83A8
293.1 B,D,F Hemagglutinin Q2RCH5 HEMA_I80A4
554.4 A,B,C,D,E,F Hemagglutinin Q03909 HEMA_I89A7
573.0 A,B,C,D,E,F Hemagglutinin P12586 HEMA_I82A6
575.1 A,B,C,D,E,F Hemagglutinin P43259 HEMA_I77A7
569.4000000000001 A,B,C,D,E,F Hemagglutinin Q2VNF2 HEMA_I78A7
568.5 A,B,C,D,E,F Hemagglutinin P03440 HEMA_I77A8
569.4000000000001 A,B,C,D,E,F Hemagglutinin Q2VND2 HEMA_I78A8
271.5 A,C,E Hemagglutinin P26141 HEMA_I84A5
571.5 A,B,C,D,E,F Hemagglutinin P11133 HEMA_I78A9
570.9 A,B,C,D,E,F Hemagglutinin P43260 HEMA_I76AC
578.4 A,B,C,D,E,F Hemagglutinin P43258 HEMA_I76AD
576.9 A,B,C,D,E,F Hemagglutinin P12582 HEMA_I77A6
571.5 A,B,C,D,E,F Hemagglutinin P26135 HEMA_I74A2
576.0 A,B,C,D,E,F Hemagglutinin P43257 HEMA_I75A2
579.6 A,B,C,D,E,F Hemagglutinin P12587 HEMA_I85A1
578.4 A,B,C,D,E,F Hemagglutinin P11134 HEMA_I82A4
571.2 A,B,C,D,E,F Hemagglutinin P26139 HEMA_I77A4
563.4 A,B,C,D,E,F Hemagglutinin Q30NQ1 HEMA_I75A0
572.1 A,B,C,D,E,F Hemagglutinin Q2RFA5 HEMA_I76A6
574.8 A,B,C,D,E,F Hemagglutinin P03435 HEMA_I75A3
572.1 A,B,C,D,E,F Hemagglutinin P26138 HEMA_I78A4
582.3 A,B,C,D,E,F Hemagglutinin P12584 HEMA_I80A7
581.7 A,B,C,D,E,F Hemagglutinin P12588 HEMA_I85A2
582.3 A,B,C,D,E,F Hemagglutinin P12585 HEMA_I82A5
577.5 A,B,C,D,E,F Hemagglutinin P26134 HEMA_I76A3
581.1 A,B,C,D,E,F Hemagglutinin P03442 HEMA_I63A3
582.0 A,B,C,D,E,F Hemagglutinin Q1PUD9 HEMA_I73A5
586.5 A,B,C,D,E,F Hemagglutinin Q2ICR0 HEMA_I72A4
586.5 A,B,C,D,E,F Hemagglutinin P03439 HEMA_I72A3
586.5 A,B,C,D,E,F Hemagglutinin P19106 HEMA_I72A2
586.5 A,B,C,D,E,F Hemagglutinin P12583 HEMA_I80A6
593.7 A,B,C,D,E,F Hemagglutinin P03449 HEMA_I71A1
594.6 A,B,C,D,E,F Hemagglutinin P03438 HEMA_I000X
594.6 A,B,C,D,E,F Hemagglutinin P03436 HEMA_I68A6
597.3 A,B,C,D,E,F Hemagglutinin Q91MA7 HEMA_I68A4
600.0 A,B,C,D,E,F Hemagglutinin P03437 HEMA_I68A0
270.9 B,D,F Hemagglutinin P17001 HEMA_I86A3
270.9 B,D,F Hemagglutinin P19699 HEMA_I86A2
270.9 B,D,F Hemagglutinin P16997 HEMA_I76AJ
284.70000000000005 B,D,F Hemagglutinin O11283 HEMA_I89A2
277.79999999999995 B,D,F Hemagglutinin Q08011 HEMA_I89A8
277.79999999999995 B,D,F Hemagglutinin Q82559 HEMA_I81A2
277.79999999999995 B,D,F Hemagglutinin P16998 HEMA_I80AB
279.29999999999995 B,D,F Hemagglutinin P16995 HEMA_I76A8
282.9 B,D,F Hemagglutinin P17000 HEMA_I71A3
284.4 B,D,F Hemagglutinin P16994 HEMA_I72A0
288.0 B,D,F Hemagglutinin P17002 HEMA_I63A4