浙江省5株H9N2亚型禽流感病毒的血凝素基因分子特征与遗传进化分析

doi:10.3969/j.issn.1004-1524.20250359

浙江农业学报 ›› 2025, Vol. 37 ›› Issue (12): 2494-2503.DOI: 10.3969/j.issn.1004-1524.20250359

浙江省5株H9N2亚型禽流感病毒的血凝素基因分子特征与遗传进化分析

刘士哲¹^,²(), 华炯钢², 陈柳², 谢荣辉³, 叶伟成², 张传亮³, 朱寅初², 冯肖肖³, 付媛², 倪征², 张存², 屈勇刚¹^,^*(), 云涛²^,^*()

1.石河子大学动物科技学院,新疆石河子 832003
2.浙江省农业科学院畜牧兽医研究所,全省畜禽生物育种重点实验室,农业农村部畜禽资源(家禽)评价利用重点实验室,家禽种业与绿色养殖技术浙江省工程研究中心,浙江杭州 310021
3.浙江省动物疫病预防控制中心,浙江杭州 311199

收稿日期:2025-05-06 出版日期:2025-12-25 发布日期:2026-01-09
作者简介:刘士哲(2000—),男,湖北荆门人,硕士研究生,主要从事畜禽疾病防控研究。E-mail:zheshiliu@163.com
通讯作者: *云涛,E-mail:yunt116@zaas.ac.cn;屈勇刚,E-mail:quyonggang@shzu.edu
基金资助:
浙江省“三农九方”科技协作计划(2023SNJF059);浙江省农业重大技术协同推广计划(2025ZDXT17-04);新疆维吾尔自治区家禽产业技术体系项目(XJARS-12-07);新疆维吾尔自治区家禽产业技术体系项目(2025xjjq-z-03)

Molecular characterization and genetic evolutionary analysis of hemagglutinin gene of five H9N2 subtype avian influenza viruses in Zhejiang Province, China

LIU Shizhe¹^,²(), HUA Jionggang², CHEN Liu², XIE Ronghui³, YE Weicheng², ZHANG Chuanliang³, ZHU Yinchu², FENG Xiaoxiao³, FU Yuan², NI Zheng², ZHANG Cun², QU Yonggang¹^,^*(), YUN Tao²^,^*()

1. College of Animal Science and Technology, Shihezi University, Shihezi 832003, Xinjiang, China
2. Zhejiang Key Laboratory of Livestock and Poultry Biotech Breeding, Key Laboratory of Livestock and Poultry Resources (Poultry) Evaluation and Utilization, Ministry of Agriculture and Rural Affairs, Zhejiang Engineering Research Center for Poultry Breeding Industry and Green Farming Technology, Institute of Animal Husbandry and Veterinary Sciences, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China
3. Zhejiang Center of Animal Disease Control and Prevention, Hangzhou 311199, China

Received:2025-05-06 Online:2025-12-25 Published:2026-01-09

摘要/Abstract

摘要：

H9N2亚型禽流感病毒(avian influenza virus, AIV)是我国禽类中最流行的低致病性禽流感(LPAIV),因其持续的抗原漂移与跨物种传播风险对养殖业和公共卫生安全构成严重威胁。为了解浙江省流行的H9N2亚型AIV的遗传进化特征,本研究对当地分离到的5株H9N2亚型AIV毒株的血凝素(HA)基因进行测序与分子特征分析。结果显示,5株分离株分属于欧亚谱系的h9.4.2.1和h9.4.2.5分支,HA基因的核苷酸和氨基酸同源性分别为83.51%~99.88%和86.33%~99.83%。值得注意的是,分离株A/Chicken/Zhejiang/HZ01/2022和A/Duck/Zhejiang/TZ01/2022与疫苗株CK/SS/94的HA基因核苷酸同源性仅为88.24%~88.35%,氨基酸同源性为89.43%~90.82%。所有分离株的HA裂解位点均呈单碱性氨基酸特征,符合低致病性病毒特性。受体结合位点分析发现存在I163T、H191N、A198T/V、Q234L和Q235M突变,这些突变可能增强病毒与人源唾液酸α-2,6-半乳糖受体的结合能力。糖基化位点分析显示,分离株存在7~8个潜在糖基化位点,其中218和551位点缺失,206和313位点增加,这些变异可能增强病毒对哺乳动物宿主的适应性。此外,HA蛋白抗原位点存在234、285和334位点突变,而181、280、281与296位点高度保守。研究表明,浙江省H9N2亚型AIV通过关键位点突变和糖基化模式改变持续进化,可能导致病毒毒力增强、疫苗保护效力下降,并增加跨物种传播风险。亟须加强本地流行毒株的分子监测,优化疫苗株匹配性,以提升防控效能并降低公共卫生威胁。

关键词: 禽流感病毒, H9N2亚型, HA基因, 遗传进化, 基因分析

Abstract:

Avian influenza virus (AIV) subtype H9N2 is the most prevalent low pathogenic avian influenza (LPAIV) in avian species in China, posing a serious threat to the farming industry and public health safety due to its persistent antigenic drift and the risk of cross-species transmission. In order to understand the genetic evolutionary characteristics of H9N2 subtype AIV prevalent in Zhejiang Province, the present study was conducted to determine the gene sequence and molecular characterization of the hemagglutinin (HA) gene of five locally isolated H9N2 subtype AIV strains. The results showed that the five isolates belonged to the h9.4.2.1 and h9.4.2.5 branches of the Eurasian lineage, and the nucleotide and amino acid homologies of the HA gene ranged from 83.51% to 99.88% and 86.33% to 99.83%, respectively. Notably, the isolates A/Chicken/Zhejiang/HZ01/2022 and A/Duck/Zhejiang/TZ01/2022 showed only 88.24%-88.35% nucleotide homology and 89.43%-90.82% amino acid homology of HA gene with the vaccine strain CK/SS/94. The HA cleavage sites of all isolates were characterized by single basic amino acid, which was consistent with the characteristics of low pathogenicity virus. Receptor binding site analysis revealed mutations I163T, H191N, A198T/V, Q234L, and Q235M, which might enhance the binding ability of the virus to the human sialic acid α-2,6-galactose receptor. Glycosylation site analysis revealed the presence of 7-8 potential glycosylation sites in the isolates, with deletions at sites 218 and 551 and additions at sites 206 and 313, and these mutations may enhance viral adaptation to mammalian hosts. In addition, HA protein antigenic sites were present with mutations at 234, 285 and 334, while sites 181, 280, 281 and 296 were highly conserved. The study suggests that Zhejiang Province H9N2 subtype AIV continues to evolve through mutations at key sites and altered glycosylation patterns, which may lead to enhanced virulence, decreased vaccine protection efficacy, and increased risk of cross-species transmission. There is an urgent need to strengthen the molecular surveillance of local endemic strains and optimize the vaccine strain match to enhance the control efficacy and reduce the public health threat.

Key words: Avian influenza virus, H9N2 subtype, HA gene, genetic evolution, genetic analysis

中图分类号:

S855.3

刘士哲, 华炯钢, 陈柳, 谢荣辉, 叶伟成, 张传亮, 朱寅初, 冯肖肖, 付媛, 倪征, 张存, 屈勇刚, 云涛. 浙江省5株H9N2亚型禽流感病毒的血凝素基因分子特征与遗传进化分析[J]. 浙江农业学报, 2025, 37(12): 2494-2503.

LIU Shizhe, HUA Jionggang, CHEN Liu, XIE Ronghui, YE Weicheng, ZHANG Chuanliang, ZHU Yinchu, FENG Xiaoxiao, FU Yuan, NI Zheng, ZHANG Cun, QU Yonggang, YUN Tao. Molecular characterization and genetic evolutionary analysis of hemagglutinin gene of five H9N2 subtype avian influenza viruses in Zhejiang Province, China[J]. Acta Agriculturae Zhejiangensis, 2025, 37(12): 2494-2503.

图/表 8

参考文献 14

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序号 Number	参考毒株 Reference strain	缩写 Abbreviation	HA基因登录号 Accession number
1	A/Turkey/Wisconsin/1/1966	TK/1/66	CY130054.1
2	A/Chicken/Shanghai/F/98	CK/F/98	AY743216.1
3	A/Chicken/Beijing/1/94	CK/1/94	AF156380.1
4	A/Chicken/Guangdong/SS/94	CK/SS/94	AF384557
5	A/Duck/Hong Kong/Y280/97	DK/Y280/97	AF156376.1
6	A/Turkey/California/189/66	TK/189/66	AF156390.1
7	A/Shorebird/Delaware/9/96	SB/9/96	AF156386.1
8	A/Chicken/Korea/SH0802/2008	CK/SH/08	HQ221677.1
9	A/Chicken/Korea/MS96-CE6/1996	CK/MS/96	GU053186.1
10	A/Chicken/Israel/178/2006	CK/178/06	EF492224.1
11	A/Chicken/Karaj/GHA/2017	CK/GHA/17	MH360730.1
12	A/Chicken/Henan/43/02	CK/43/02	DQ064369.1
13	A/Goose/China/JY100405/2018	GS/JY/18	OQ804402.1
14	A/Chicken/Jiangsu/J2303/2018	CK/J2303/18	MK553246.1
15	A/Chicken/Rizhao/1436/2013	CK/1436/13	KF259177.1
16	A/Chicken/China/GD2021/2021	CK/GD2021/21	MW548848.1
17	A/Chicken/Daye/DY0602/2017	CK/DY0602/17	MF794999.1
18	A/Chicken/China/355/2017	CK/355/17	MN385379.1
19	A/Chicken/Guangzhou/18/2019	CK/18/19	OQ054042.1
20	A/Chicken/China/D3/2022	CK/D3/22	OR528233.1
21	A/Duck/Guangdong/FS91/2022	DK/FS91/22	OQ826094.1
22	A/Environment/Xiamen/01/2021	EN/01/21	ON856628.1
23	A/Chicken/Shandong/6/96	CK/6/96	DQ064376.1
24	A/Chicken/Shandong/1/2000	CK/1/00	FJ793404.1
25	A/Chicken/Jilin/A/2012	CK/A/12	KF886521.1
26	A/Chicken/Jiangxi/X1459/2018	CK/X1459/18	MK552958.1
27	A/Chicken/Guizhou/B11/2011	CK/B11/11	JN804200.1
28	A/Chicken/Guangdong/WBQ02/2012	CK/WB/12	KJ768997.1

序号 Number	参考毒株 Reference strain	缩写 Abbreviation	HA基因登录号 Accession number
1	A/Turkey/Wisconsin/1/1966	TK/1/66	CY130054.1
2	A/Chicken/Shanghai/F/98	CK/F/98	AY743216.1
3	A/Chicken/Beijing/1/94	CK/1/94	AF156380.1
4	A/Chicken/Guangdong/SS/94	CK/SS/94	AF384557
5	A/Duck/Hong Kong/Y280/97	DK/Y280/97	AF156376.1
6	A/Turkey/California/189/66	TK/189/66	AF156390.1
7	A/Shorebird/Delaware/9/96	SB/9/96	AF156386.1
8	A/Chicken/Korea/SH0802/2008	CK/SH/08	HQ221677.1
9	A/Chicken/Korea/MS96-CE6/1996	CK/MS/96	GU053186.1
10	A/Chicken/Israel/178/2006	CK/178/06	EF492224.1
11	A/Chicken/Karaj/GHA/2017	CK/GHA/17	MH360730.1
12	A/Chicken/Henan/43/02	CK/43/02	DQ064369.1
13	A/Goose/China/JY100405/2018	GS/JY/18	OQ804402.1
14	A/Chicken/Jiangsu/J2303/2018	CK/J2303/18	MK553246.1
15	A/Chicken/Rizhao/1436/2013	CK/1436/13	KF259177.1
16	A/Chicken/China/GD2021/2021	CK/GD2021/21	MW548848.1
17	A/Chicken/Daye/DY0602/2017	CK/DY0602/17	MF794999.1
18	A/Chicken/China/355/2017	CK/355/17	MN385379.1
19	A/Chicken/Guangzhou/18/2019	CK/18/19	OQ054042.1
20	A/Chicken/China/D3/2022	CK/D3/22	OR528233.1
21	A/Duck/Guangdong/FS91/2022	DK/FS91/22	OQ826094.1
22	A/Environment/Xiamen/01/2021	EN/01/21	ON856628.1
23	A/Chicken/Shandong/6/96	CK/6/96	DQ064376.1
24	A/Chicken/Shandong/1/2000	CK/1/00	FJ793404.1
25	A/Chicken/Jilin/A/2012	CK/A/12	KF886521.1
26	A/Chicken/Jiangxi/X1459/2018	CK/X1459/18	MK552958.1
27	A/Chicken/Guizhou/B11/2011	CK/B11/11	JN804200.1
28	A/Chicken/Guangdong/WBQ02/2012	CK/WB/12	KJ768997.1

毒株 Strain	分离时间 Time	地区 Area	宿主 Specie	基因 Gene	同源性最高的病毒株 Viral strain with the highest homology	核酸同源性 Nucleotide homology/%	氨基酸同源性 Amino acid homology/%	登录号 Accession number
A/Chicken/Zhejiang/ HZ01/2022 (CK/01/22)	2022	杭州 Hangzhou	鸡 Gallus demesticus	HA	A/Chicken/Jiangxi/ X1459/2018(H9N2)	98.75	99.29	MK552958.1
A/Quails/Zhejiang/ SY01/2012 (QU/01/12)	2012	上虞 Shangyu	鹌鹑 Coturnix japonica	HA	A/Chicken/Shanghai/ S2277/2017(H9N2)	98.70	97.39	MK554249.1
A/Quails/Zhejiang/ JX535/2012 (QU/535/12)	2012	嘉兴 Jiaxing	鹌鹑 Coturnix japonica	HA	A/Chicken/Shanghai/ S2277/2017(H9N2)	98.88	98.32	MK554249.1
A/Chicken/Zhejiang/ XS541/2018 (CK/541/18)	2018	象山 Xiangshan	鸡 Gallus demesticus	HA	A/Chicken/Jilin/ A/2012(H9N2)	99.81	99.44	KF886521.1
A/Duck/Zhejiang/ TZ01/2022 (DK/01/22)	2022	台州 Taizhu	鸭 Anas platyrhynchos	HA	A/Chicken/Jiangxi/ X1459/2018(H9N2)	99.75	99.29	MK552958.1

毒株 Strain	分离时间 Time	地区 Area	宿主 Specie	基因 Gene	同源性最高的病毒株 Viral strain with the highest homology	核酸同源性 Nucleotide homology/%	氨基酸同源性 Amino acid homology/%	登录号 Accession number
A/Chicken/Zhejiang/ HZ01/2022 (CK/01/22)	2022	杭州 Hangzhou	鸡 Gallus demesticus	HA	A/Chicken/Jiangxi/ X1459/2018(H9N2)	98.75	99.29	MK552958.1
A/Quails/Zhejiang/ SY01/2012 (QU/01/12)	2012	上虞 Shangyu	鹌鹑 Coturnix japonica	HA	A/Chicken/Shanghai/ S2277/2017(H9N2)	98.70	97.39	MK554249.1
A/Quails/Zhejiang/ JX535/2012 (QU/535/12)	2012	嘉兴 Jiaxing	鹌鹑 Coturnix japonica	HA	A/Chicken/Shanghai/ S2277/2017(H9N2)	98.88	98.32	MK554249.1
A/Chicken/Zhejiang/ XS541/2018 (CK/541/18)	2018	象山 Xiangshan	鸡 Gallus demesticus	HA	A/Chicken/Jilin/ A/2012(H9N2)	99.81	99.44	KF886521.1
A/Duck/Zhejiang/ TZ01/2022 (DK/01/22)	2022	台州 Taizhu	鸭 Anas platyrhynchos	HA	A/Chicken/Jiangxi/ X1459/2018(H9N2)	99.75	99.29	MK552958.1

毒株 Strain	核苷酸同源性Nucleotide homology				氨基酸同源性Amino acid homology
毒株 Strain	CK/SS/94	CK/F/98	CK/6/96	CK/1/00	CK/SS/94	CK/F/98	CK/6/96	CK/1/00
CK/01/22	88.35	87.88	87.70	87.44	90.18	90.18	89.64	89.62
QU/01/12	87.84	86.60	86.78	86.78	89.70	88.39	88.80	88.53
QU/535/12	88.46	87.21	88.08	87.41	90.82	89.14	89.57	89.83
CK/541/18	96.12	94.67	95.49	94.88	97.56	96.80	96.43	96.35
DK/01/22	88.24	87.76	87.58	87.31	90.00	90.00	89.46	89.43

浙江省5株H9N2亚型禽流感病毒的血凝素基因分子特征与遗传进化分析

Molecular characterization and genetic evolutionary analysis of hemagglutinin gene of five H9N2 subtype avian influenza viruses in Zhejiang Province, China

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分离株 Strain	裂解位点 Cleavage site	左侧臂 Left side wall	右侧臂 Right side wall	受体结合位点 Receptor binding site
分离株 Strain	333-341	232-237	146-150	109	161	163	191	198	202	203	224
CK/GHA/17*	PARSSR↓GLF	NGLIGR	GTSKS	Y	W	T	H	A	L	Y	L
CK/BJ/94*	PARSSR↓GLF	NGQQGR	GTSKA	Y	W	T	N	V	L	Y	Q
CK/F/98*	PARSSR↓GLF	NGQQGR	GTSKA	Y	W	T	N	A	L	Y	Q
CK/SS/94*	PAGSSR↓GLF	NGQQGR	GTSKA	Y	W	T	N	A	L	Y	V
DK/Y280/97*	PARSSR↓GLF	NGLQGR	GTSKA	Y	W	T	N	T	L	Y	L
CK/43/02*	PARSSR↓GLF	NGLQGR	GTSRA	Y	W	T	N	A	L	Y	L
EN/01/2021*	PSRSSR↓GLF	NGLMGR	GTSNT	Y	W	N	N	V	L	Y	L
CK/D3/2022*	PSRSSR↓GLF	NGLMGR	GTSKT	Y	W	N	N	V	L	Y	L
CK/01/22	PSRSSR↓GLF	NGLMGR	GTSTA	Y	W	T	N	T	L	Y	L
QU/01/12	PARSSR↓GLF	NGQQGR	GTSIS	Y	W	T	H	A	L	Y	V
QU/535/12	PARSSR↓GLF	NGQQGR	GTSIS	Y	W	T	H	T	L	Y	V
CK/541/18	PARLSR↓GLF	NGQQGR	GTSKA	Y	W	T	N	T	L	Y	L
DK/01/22	PSRSSR↓GLF	NGLMGR	GTSTA	Y	W	T	N	T	L	Y	L

分离株 Strain	糖基化位点Glycosylation site
分离株 Strain	29	82	141	206	218	298	305	313	492	551
CK/GHA/17*	NSTE	NPSC	NVTY			NSTL	NISK		NGTY	NGSC
CK/BJ/94*	NSTE	NPSC	NVTY		NRTF	NTTL	NVSK		NGTY	NGSC
CK/F/98*	NSTE	NPSC	NVSY			NTTL	NVSK		NGTY	NGSC
CK/SS/94*	NSTE	NPSC	NVSY		NRTF	NTTL	NVSK		NGTY	NGSC
Dk/Y280/97*	NSTE	NPSC	NVSY		NRTF	NTTL	NVSK		NGTY
CK/43/02*	NSTE	NPSC	NVSY		NRTF	NTTL	NVSK		NGTY	NGSC
EN/01/2021*	NSTE	NPSC	NVSY			NTTL	NVSK	NCSK	NGTY	NGSC
CK/D3/2022*	NSTE	NPSC	NVSY			NTTL	NVSK	NCSK	NGTY	NGSC
CK/01/22	NSTE	NPSC	NVSY			NTTL	NVSK	NCSK	NGTY	NGSC
QU/01/12	NSTE	NPSC	NVSY	NDTT	NRTF	NSTL	NISK		NGTY
QU/535/12	NSTE	NPSC	NVSY	NDTT	NRTF	NSTL	NISK		NGTY
CK/541/18	NSTE	NPSC	NVSY		NRTF	NTTL	NVSK		NGTY
DK/01/22	NSTE	NPSC	NVSY			NTTL	NVSK	NCSK	NGTY	NGSC

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