微生物学报

Virus name	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18
1	100	96.8	97.4	97.4	92.7	97.3	97.5	97.6	97.1	96.9	97.3	93.1	93.2	91.9	92.1	93.1	92.9	92.3
2		100	96.1	96.1	94.6	96.1	96.2	96.4	95.9	95.5	96.0	94.9	95.0	93.4	93.8	95.0	94.6	94.1
3			100	99.3	92.3	97.9	99.3	98.3	99.1	98.8	99.2	92.7	92.7	91.5	91.8	92.6	92.4	92.0
4				100	92.3	98.0	99.2	98.4	99.5	99.5	99.6	92.7	92.8	91.4	91.7	92.6	92.4	92.1
5					100	92.1	92.2	92.3	91.9	91.7	92.2	93.3	93.3	92.0	92.4	93.1	92.9	92.5
6						100	98.1	99.4	97.8	97.5	98.2	92.6	92.7	91.4	91.7	92.5	92.3	91.8
7							100	98.6	99.1	98.8	99.2	92.8	92.8	91.5	91.8	92.6	92.4	92.1
8								100	98.2	98.0	98.3	92.9	93.0	91.7	92.0	92.8	92.5	92.1
9									100	99.0	99.5	92.5	92.5	91.2	91.5	92.3	92.1	91.8
10										100	99.2	92.2	92.3	91.0	91.3	92.1	91.9	91.6
11											100	92.7	92.7	91.4	91.7	92.6	92.4	92.0
12												100	99.9	96.9	97.3	93.5	93.2	92.8
13													100	96.9	97.3	93.5	93.2	92.8
14														100	98.9	92.4	92.0	91.6
15															100	92.7	92.3	91.9
16																100	98.2	94.5
17																	100	94.1
18																		100

Virus name	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18
1	100	96.8	97.4	97.4	92.7	97.3	97.5	97.6	97.1	96.9	97.3	93.1	93.2	91.9	92.1	93.1	92.9	92.3
2		100	96.1	96.1	94.6	96.1	96.2	96.4	95.9	95.5	96.0	94.9	95.0	93.4	93.8	95.0	94.6	94.1
3			100	99.3	92.3	97.9	99.3	98.3	99.1	98.8	99.2	92.7	92.7	91.5	91.8	92.6	92.4	92.0
4				100	92.3	98.0	99.2	98.4	99.5	99.5	99.6	92.7	92.8	91.4	91.7	92.6	92.4	92.1
5					100	92.1	92.2	92.3	91.9	91.7	92.2	93.3	93.3	92.0	92.4	93.1	92.9	92.5
6						100	98.1	99.4	97.8	97.5	98.2	92.6	92.7	91.4	91.7	92.5	92.3	91.8
7							100	98.6	99.1	98.8	99.2	92.8	92.8	91.5	91.8	92.6	92.4	92.1
8								100	98.2	98.0	98.3	92.9	93.0	91.7	92.0	92.8	92.5	92.1
9									100	99.0	99.5	92.5	92.5	91.2	91.5	92.3	92.1	91.8
10										100	99.2	92.2	92.3	91.0	91.3	92.1	91.9	91.6
11											100	92.7	92.7	91.4	91.7	92.6	92.4	92.0
12												100	99.9	96.9	97.3	93.5	93.2	92.8
13													100	96.9	97.3	93.5	93.2	92.8
14														100	98.9	92.4	92.0	91.6
15															100	92.7	92.3	91.9
16																100	98.2	94.5
17																	100	94.1
18																		100

RNA type	RNA name	Targeting molecule	Function mode	References
miRNA	miR-217	TGF-b-activated kinase 1 (TAK1)	Inhibition of protein expression (IPE)	[38]
	miR-217-5p	Nucleotide oligomerization domain 1 (NOD1)	IPE	[39]
	miR-3570	Mitochondrial antiviral signaling protein (MAVS)	IPE	[40]
	miR-122	MAVS	IPE	[41]
	miR-2187	Tumor necrosis factor receptor related factor 6 (TRAF6)	IPE	[42]
	miR-2187-3p	TANK-binding kinase 1 (TBK1)	IPE	[43]
	miR-181b-2	TIR-domain-containing adapter-inducing interferon-b (TRIF)	IPE	[44]
	miR-21-1	TRIF	IPE	[44]
	miR-210	Stimulator of IFN genes (STING)	IPE	[45]
	miR-214	N and P genes of SCRV	Inhibition of viral replication	[46]
lncRNA	MIR2187HG	miR-2187-3p	Pre-miRNA	[43]
	MIR122HG	miR-122, miR-122-5p	Pre-miRNA	[47-48]
	lncRNA AANCR	miR-210	Sponge effect	[49]
	lncRNA MARL	miR-122	Sponge effect	[41]
	lncRNA NARL	miR-217-5p	Sponge effect	[39]
circRNA	circRasGEF1B	miR-21-3p (targeting MITA)	Sponge effect	[50]
	circCBL	miR-125a-1-3p (targeting MITA)	Sponge effect	[51]
	circDtx1	miR-15a-5p (targeting TRIF)	Sponge effect	[52]
	circSamd4a	miR-29a-3p (targeting STING)	Sponge effect	[53]
	circRNF217	miR-130-3p (targeting NOD1)	Sponge effect	[54]
	circBCL2L1	miR-30c-3-3p (targeting TRAF6)	Sponge effect	[55]
	circPIKfyve	miR-21-3p (targeting MAVS)	Sponge effect	[56]
	circVPS13D	VPS13D-170aa (targeting MAVS)	Coding protein	[57]
	circMORC3	MORC3-84aa (targeting TRIF)	Coding protein	[58]
	circYthdc2	Ythdc2-170aa (targeting STING)	Coding protein	[59]
	circNLRP12	NLRP12-119aa (targeting SCRV)	Coding protein	[60]

RNA type	RNA name	Targeting molecule	Function mode	References
miRNA	miR-217	TGF-b-activated kinase 1 (TAK1)	Inhibition of protein expression (IPE)	[38]
	miR-217-5p	Nucleotide oligomerization domain 1 (NOD1)	IPE	[39]
	miR-3570	Mitochondrial antiviral signaling protein (MAVS)	IPE	[40]
	miR-122	MAVS	IPE	[41]
	miR-2187	Tumor necrosis factor receptor related factor 6 (TRAF6)	IPE	[42]
	miR-2187-3p	TANK-binding kinase 1 (TBK1)	IPE	[43]
	miR-181b-2	TIR-domain-containing adapter-inducing interferon-b (TRIF)	IPE	[44]
	miR-21-1	TRIF	IPE	[44]
	miR-210	Stimulator of IFN genes (STING)	IPE	[45]
	miR-214	N and P genes of SCRV	Inhibition of viral replication	[46]
lncRNA	MIR2187HG	miR-2187-3p	Pre-miRNA	[43]
	MIR122HG	miR-122, miR-122-5p	Pre-miRNA	[47-48]
	lncRNA AANCR	miR-210	Sponge effect	[49]
	lncRNA MARL	miR-122	Sponge effect	[41]
	lncRNA NARL	miR-217-5p	Sponge effect	[39]
circRNA	circRasGEF1B	miR-21-3p (targeting MITA)	Sponge effect	[50]
	circCBL	miR-125a-1-3p (targeting MITA)	Sponge effect	[51]
	circDtx1	miR-15a-5p (targeting TRIF)	Sponge effect	[52]
	circSamd4a	miR-29a-3p (targeting STING)	Sponge effect	[53]
	circRNF217	miR-130-3p (targeting NOD1)	Sponge effect	[54]
	circBCL2L1	miR-30c-3-3p (targeting TRAF6)	Sponge effect	[55]
	circPIKfyve	miR-21-3p (targeting MAVS)	Sponge effect	[56]
	circVPS13D	VPS13D-170aa (targeting MAVS)	Coding protein	[57]
	circMORC3	MORC3-84aa (targeting TRIF)	Coding protein	[58]
	circYthdc2	Ythdc2-170aa (targeting STING)	Coding protein	[59]
	circNLRP12	NLRP12-119aa (targeting SCRV)	Coding protein	[60]

鳜鱼弹状病毒研究进展

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刘晓瑜 , 张建华 , 朱晓薇 , 魏永伟 ^*

微生物学报 | 综述 2026,66(1): 134-148

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微生物学报 | 综述 2026, 66(1): 134-148

鳜鱼弹状病毒研究进展

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刘晓瑜, 张建华, 朱晓薇, 魏永伟^*

作者信息

绍兴文理学院医学院，浙江绍兴

Research progress in Siniperca chuatsi rhabdovirus

Xiaoyu LIU, Jianhua ZHANG, Xiaowei ZHU, Yongwei WEI^*

Affiliations

School of Medicine, Shaoxing University, Shaoxing, Zhejiang, China

出版时间: 2026-01-04 doi: 10.13343/j.cnki.wsxb.20250576

文章导航

摘要

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鳜鱼是亚洲国家极为重要的经济养殖鱼类之一。随着人工养殖规模不断扩大，传染病的发生成为鳜鱼养殖业的重要危害因素，也是鳜鱼养殖产业发展的突出问题。鳜鱼弹状病毒(Siniperca chuatsi rhabdovirus, SCRV)是感染鳜鱼的一种重要病原体。近几年，SCRV的研究取得了较大进展，目前尚无相关综述文章。本文就SCRV的病毒特性、病毒拯救、与宿主互作关系及防控策略等方面的研究进展进行综述与讨论，并对面临的问题展开分析。

关键词

鳜鱼弹状病毒 / 病毒特性 / 病毒拯救 / 宿主互作 / 防控策略

Abstract

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The mandarin fish (Siniperca chuatsi) is one of the most economically important cultured fish species in Asian countries. With the expansion of artificial farming, infectious diseases have become a major threat to the mandarin fish farming industry, posing a challenge to its sustainable development. Siniperca chuatsi rhabdovirus (SCRV) is a major pathogen infecting this fish species. In recent years, substantial progress has been made in the research on SCRV, yet no comprehensive review is currently available. This paper summarizes and discusses the research advances in SCRV, including viral characteristics, virus rescue, host-virus interactions, and prevention strategies, while also analyzing the current challenges in this field.

Key words

Siniperca chuatsi rhabdovirus (SCRV) / viral characteristics / virus rescue / host-virus interaction / prevention strategies

引用本文

刘晓瑜, 张建华, 朱晓薇, 魏永伟. 鳜鱼弹状病毒研究进展. 微生物学报, 2026 , 66 (1) : 134 -148 . DOI: 10.13343/j.cnki.wsxb.20250576

Xiaoyu LIU, Jianhua ZHANG, Xiaowei ZHU, Yongwei WEI. Research progress in Siniperca chuatsi rhabdovirus[J]. Acta Microbiologica Sinica, 2026 , 66 (1) : 134 -148 . DOI: 10.13343/j.cnki.wsxb.20250576

正文

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鳜鱼因其肉质鲜美、营养丰富且无肌间刺等特点深受消费者喜爱。鳜鱼生长迅速，市场价值高，是亚洲国家极为重要的经济养殖鱼类之一。中国鳜鱼养殖年产量达30多万t，产值超过200亿元；然而，随着鳜鱼人工养殖规模扩大以及集约化养殖的推广应用，传染病的发生成为鳜鱼养殖业的重要危害，造成巨大经济损失，已成为阻碍鳜鱼可持续养殖的关键因素，是鳜鱼养殖产业发展中最为突出的问题之一^[1]。

弹状病毒是水生动物新发与再发病毒中较为常见的病毒种类之一^[2]。鳜鱼弹状病毒(Siniperca chuatsi rhabdovirus, SCRV)是感染鳜鱼的重要病原体。SCRV最早在被感染的鳜鱼组织中被观察到；发病鳜鱼的临床表现为口腔周围、鳍基部和尾部充血，部分病鱼眼球突出；病死鳜鱼的鳃丝、鳍条基部、肝脏和脾脏有出血点，肾脏肿大且出血^[3-4]。SCRV感染鳜鱼2 d后即可导致其死亡，4 d内死亡率可达70%-100%^[5]。除鳜鱼外，SCRV还可感染鲈鱼，发病鲈鱼表现为螺旋形游泳行为，身体弯曲，眼球突出，腹部膨胀，下颌至腹部有瘀点出血，肝脏肿胀且出血；养殖场暴发该病后死亡率达50%，实验室感染死亡率达85%^[4,6]。

自Tao等^[7-9]克隆出第一株SCRV全基因组后，近几年SCRV的研究取得了较大进展。例如，GenBank数据库中提交的全基因组序列数量显著增加，反向遗传系统得以建立，在感染后宿主转录组及非编码RNA分析、病毒稳定性等方面的研究也取得了很大进步。目前，国内外尚无关于SCRV的综述文章。本文就SCRV的病毒特性、病毒拯救、与宿主互作关系和防控策略等方面的研究进行综述和讨论，并对研究面临的问题进行分析，以期加深对SCRV的认识和理解，为SCRV的未来研究提供借鉴与参考。

1 病毒学特性

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1.1 SCRV的分类地位

弹状病毒科(Rhabdoviridae)包含56个病毒属，其中Perhabdovirus、Siniperhavirus、Scophrhavirus、Sprivivirus和诺拉弹状病毒属(Novirhabdovirus)这5个属的成员可感染鱼类。SCRV是Siniperhavirus的成员^[10]。此前，SCRV被归类于Perhabdovirus病毒属，在国际病毒分类委员会(International Committee on Taxonomy of Viruses, ICTV)最新的病毒分类报告中它被划归到一个新的病毒属——中国鲈鱼病毒属(Siniperhavirus)^[10]。由于Siniperhavirus是新分类的病毒属，尚无对应的中文名称，该属中文名为本文暂定名。从大口黑鲈、乌鳢和黄鳝中分离出的弹状病毒分别被称为大口黑鲈弹状病毒(micropterus salmoides rhabdovirus, MSRV)^[11-13]、杂交鳢弹状病毒(hybrid snakehead rhabdovirus, HSHRV)^[14-15]或乌鳢水泡病毒(snakehead fish vesiculovirus, SHVV)^[16]、黄鳝弹状病毒(Chinese rice-field eel rhabdovirus, CrERV)^[17-18]或传染性出血综合征病毒(infectious haemorrhagic syndrome virus, IHSV)^[19]，这些病毒在进化关系上较为接近^[20]，基因组一致性达90%以上(详见下文分析)。

1.2 SCRV基因组及其病毒粒子的分子结构

SCRV于1999年首次从感染的鳜鱼中分离得到^[3-4]，其基因组为单股负链RNA，大小为11 545 bp，基因组注册号为DQ399789^[6-7]。如图1所示，SCRV基因组编码5种结构蛋白，分别为核蛋白(nucleoprotein, N)、磷蛋白(phosphoprotein, P)、基质蛋白(matrix protein, M)、糖蛋白(glycoprotein, G)和RNA聚合酶(RNA polymerase, L)。基因组从3′至5′方向依次为3′前导序列(3′ leader)-N-P-M-G-L-5′非编码区(5′ trailer)；SCRV病毒粒子长约100-430 nm，直径45-100 nm，具有典型的子弹状形态，且有囊膜结构；病毒粒子内部的核酸由N蛋白包裹，其表面是由G蛋白构成的纤突状突起，负责病毒的吸附和进入；病毒粒子内表面由基质蛋白M组成；核蛋白N与磷蛋白P和RNA聚合酶蛋白L结合共同组成RNA依赖的RNA聚合酶复合体，负责病毒基因组RNA和病毒蛋白mRNA的合成^[7,21]。

1.3 SCRV基因组序列特征和进化关系

截至2025年9月，在GenBank数据库中提交的具有全基因组序列的SCRV毒株共有5株。为分析SCRV基因组序列特征和进化关系，从GenBank数据库中下载中国鲈鱼病毒属中所有具有全基因组的病毒株序列，应用DNASTAR 5.0中的Clustal NJ进行多序列比对，随后开展全基因组序列同源性分析。如表1所示，各SCRV毒株之间的全基因组序列一致性为92.3%-99.3%，SCRV与MSRV之间的一致性为91.9%-99.6%。SCRV与分离自乌鳢的弹状病毒的一致性为91.4%-95.0%，与分离自黄鳝的弹状病毒一致性为92.0%-95.0%。值得指出的是，SCRV的SS株(GenBank登录号为PP784252)与其他所有毒株的同源性均较低(SS株与其他所有毒株的一致性为91.7%-94.6%)。若不计SS株，各SCRV毒株之间的全基因组一致性为96.1%-99.3%，SCRV与MSRV之间的一致性为95.5%-99.6%。发育树分析显示，分离自鳜鱼的SCRV和分离自鲈鱼的MSRV聚为一个大的分支(SS株除外)，分离自乌鳢的弹状病毒和分离自黄鳝的弹状病毒分别聚类在一起，而分离自鳜鱼的SS株则独自形成一个进化分支，并且在进化关系上SS株与分离自乌鳢和黄鳝的弹状病毒更为接近(图2)。上述结果表明：(1) SCRV和MSRV之间的同源性与SCRV和MSRV各自内部之间的同源性相近，SCRV和MSRV在进化关系上可能属于同种病毒；(2) SCRV的SS株是不同于其他毒株的独立进化分支。

2 SCRV反向遗传系统的建立

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经典遗传学是从生物的表型到遗传物质来研究生命现象。反向遗传学则是由生物基因组序列重新装配出具有生命活性的个体。由于病毒基因组相对较小，易于操作，反向遗传学成为病毒研究的重要领域。病毒反向遗传系统也被称为“病毒拯救”。已鉴定的水生动物RNA病毒有上百种之多，但建立的水生动物RNA病毒反向遗传系统却鲜有报道。在弹状病毒科中，水生动物弹状病毒仅有诺拉弹状病毒属的传染性造血器官坏死病毒(infectious hematopoietic necrosis virus, IHNV)^[22]、病毒性出血败血症病毒(viral hemorrhagic septicemia virus, VHSV)^[23]、乌鳢弹状病毒(snakehead rhabdovirus, SHRV)^[24]和牙鲆弹状病毒(hirame rhabdovirus, HIRRV)^[25]；鲤春弹状病毒属的鲤春病毒血症病毒(spring viraemia of carp virus, SVCV)^[26]以及中国鲈鱼病毒属的SHVV^[27]和SCRV^[28]建立了反向遗传系统。SCRV的反向遗传系统由魏永伟团队建立，该团队利用克隆的SCRV S3株作为母源毒株，成功建立了带有分子标记和荧光报告基因的重组株^[29]。这为SCRV基因功能研究、药物筛选、疫苗研制和开发病毒载体提供了有力工具，具有广阔的应用前景。

3 SCRV与宿主的关系

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3.1 SCRV的感染宿主范围

已发现的SCRV可感染的自然宿主仅有鳜鱼和大口黑鲈。在实验室条件下，SCRV可感染斑马鱼(Danio rerio)^[30]和鮸鱼(Miichthys miiuy)^[31]。体外细胞感染实验表明，SCRV除了能感染来源于自然宿主鳜鱼和大口黑鲈的细胞株外，还可感染其他多种鱼源细胞，如鲤鱼上皮细胞(epithelioma papulosum cyprini cells, EPC cells)^[30]、草鱼鳍细胞(grass carp fins cells, GCF cells)^[5]、胖头鲤细胞(fathead minnow cells, FHM cells)^[9]、真鲷脑细胞(red sea bream brain cells, RSBB cells)^[32]和E11细胞(条纹蛇头鱼SSN-1细胞(striped snakehead cells, SSN-1 cells)的一个克隆株)^[33]。此外，SCRV在28 ℃也可感染哺乳动物细胞Vero E6 (非洲绿毛猴肾细胞)以及人源细胞5637 (膀胱癌细胞)、BGC-823 (胃腺癌细胞)和HCC1937 (乳腺癌细胞)^[30]。细胞感染实验说明SCRV的受体广泛分布于不同物种。由此推测，在自然界中除了已知的自然宿主鳜鱼和大口黑鲈外，其他鱼类很可能也能感染SCRV或成为SCRV的潜在携带者。

3.2 SCRV感染宿主情况

通过反向遗传系统拯救的携带报告基因的rS3-LeNEGFP实现了SCRV感染可视化。利用rS3-LeNEGFP在斑马鱼体内的动态示踪表明，SCRV主要侵入斑马鱼的肝、心和脑，在感染后24 h可在肝和心检测到大量荧光蛋白的表达，且这种表达一直持续到感染后的第10天；在心脏的心房和心室均观察到大面积且很强的荧光信号；在感染后第3天可在脑中检测到荧光蛋白的表达；斑马鱼感染SCRV后的重要临床症状表现为不规则游动、打转或快速兴奋游动，这些症状可能与SCRV感染脑有直接关系；此外，在感染的雄性斑马鱼精原细胞中检测到了荧光信号，那么SCRV能否通过精子进行垂直传播值得进一步研究^[29]。

3.3 SCRV感染后的宿主在分子水平的反应

SCRV感染宿主后在分子水平的反应研究得益于现代测序技术的进步。相关研究主要集中在mRNA转录组分析、小RNA (microRNA, miRNA)、长链非编码RNA (long noncoding RNA, lncRNA)和环状RNA (circular RNA, circRNA) 4个方面。

SCRV感染鳜鱼后的肾脏转录组分析显示差异表达基因(differentially expressed genes, DEGs)主要涉及免疫反应和细胞凋亡等方面，以及RIG-I样受体/Toll样受体/NOD样受体/C型凝集素受体信号通路、p53通路和代谢通路等；感染后干扰素-I (interferon I, IFN-I)、白细胞介素8 (interleukin 8, IL8)、干扰素调节因子3 (interferon regulatory factor 3, IRF3)、melanoma differentiation-associated protein 5 (MDA5)和laboratory of genetics and physiology 2 (LGP2)的表达量显著上调^[34]。在体外细胞水平，SCRV感染E11细胞后的DEGs差异主要涉及丝裂原活化蛋白激酶(mitogen-activated protein kinase, MAPK)信号通路、phosphatidylinositol 3-kinase-protein kinase B (PI3K-Akt)信号通路、内吞作用和凋亡等^[33]。感染SCSC细胞(鳜鱼皮肤细胞)后DEGs响应的关键差异为“细胞因子-细胞因子受体相互作用”和“干扰素相关通路”，其中5种干扰素刺激基因(IFI4407、IFI35、Viperin、IFIT1和IFIT5)的表达量显著上调^[35]。显然，上述动物水平和不同细胞转录组分析结果并不一致，说明不同宿主细胞以及在细胞水平和动物水平受到SCRV感染后其反应存在较大差异，引起这种差异的分子机制有待研究。

miRNA是一类长度约22个核苷酸的非编码RNA，通过调控靶基因的表达参与多种生物学调控过程。miRNA可与负责编码蛋白质的mRNA的3′非编码区结合，引起mRNA翻译抑制或诱导其降解^[36-37]。用SCRV感染鮸鱼的动物模型研究表明，多种miRNA可通过抑制先天免疫过程中核因子κB (nuclear factor kappa-B, NF-κB)和IRF3信号通路上游某个关键蛋白的表达，进而抑制炎症因子和抗病毒因子的表达(表2)。lncRNA是一类长度约200个核苷酸且不编码蛋白质的RNA分子，在转录调控、翻译、表观遗传修饰等生物学过程中发挥着重要作用。lncRNA在病毒感染后的先天免疫反应中起着重要的调控作用^[61-62]。lncRNA一方面可作为miRNA的前体，经剪切加工成成熟的miRNA发挥调控作用；另一方面，lncRNA作为“miRNA分子海绵”竞争性地吸附miRNA，通过调控miRNA间接调控先天免疫反应过程。SCRV感染宿主后lncRNA也可通过上述2种方式参与调控先天免疫反应过程(表2)。circRNA是一类具有共价闭合环状结构的RNA，其独特的结构赋予其抗核酸外切酶降解的能力，因而具有高稳定性^[63-65]。研究发现SCRV感染后的circRNA调控主要表现在以下途径：(1) 如同上述的lncRNA，circRNA作为miRNA分子海绵吸附miRNA，解除miRNA对NF-κB/IRF3信号通路上游某个特定蛋白的抑制，进而增强先天抗病毒免疫反应；(2) circRNA通过编码蛋白质发挥调控作用，所编码的蛋白质靶向NF-κB/IRF3信号通路上游的特定蛋白以降低抗病毒细胞因子表达，或直接抑制SCRV复制。当前已鉴定的在SCRV感染中发挥调控作用的circRNA及其靶向分子如表2所示。

如图3所示，SCRV感染后miRNA、lncRNA和circRNA通过lncRNA-miRNA-mRNA-X蛋白轴、circRNA-miRNA-mRNA-X蛋白轴和circRNA-编码蛋白-X蛋白(或病毒)轴3种基本模式发挥宿主细胞的抗病毒调控作用。然而，值得指出的是，由于已有的研究均是孤立进行的，上述3种路径哪种占主导，或者3种路径之间是否相互影响还有待进一步系统研究。在SCRV引起的宿主抗病毒免疫调控中，其核心是IRF3/NF-κB信号通路上游的某个关键蛋白(X蛋白)受到影响，进而引起下游炎症因子和抗病毒因子表达受到调控。由上述已报道的相关研究可知，同种X蛋白(或其mRNA)可作为不同调控因子的靶标，如miR-3570、miR-122、miR-21-3p和VPS13D-170aa均可靶向MAVS，miR-181b-2、miR-21-1、miR-15a-5p和MORC3-84aa均可靶向TRIF。这种基于同种X蛋白的不同调控路径中哪种miRNA调控占主导作用，或者不同miRNA靶向同种靶标之间有何关系及相互影响如何均有待进一步研究。

4 SCRV检测技术

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当前建立的可用于临床样品检测SCRV的方法有普通PCR和基于荧光探针的TaqMan实时荧光定量PCR (quantitative real-time PCR, qPCR) 2种。普通PCR是与感染鳜鱼的另外2种重要病毒传染性脾肾坏死病毒(infectious spleen and kidney necrosis virus, ISKNV)和鳜鱼蛙病毒(Siniperca chuatsi ranairidovirus, SCRIV)联合检测，分别是SCRV和ISKNV双重PCR^[66]、SCRV和SCRIV双重PCR^[67]以及SCRV、ISKNV和SCRIV三重PCR^[68]。单独检测SCRV的TaqMan qPCR灵敏度高于普通PCR^[69]。除上述用于临床样本检测的方法外，Lin等^[70]建立了基于荧光染料的实时荧光定量逆转录PCR (real-time RT-PCR, RT-qPCR)，用于区分SCRV感染过程中产生的病毒基因组RNA、病毒基因组互补RNA和mRNA。Niu等^[71]用纯化的SCRV病毒颗粒免疫BALB/c小鼠，再通过细胞融合获得2株单克隆抗体4H8和4E12，这2株抗体能特异性识别SCRV的G蛋白；利用这2株单克隆抗体建立了双抗体夹心酶联免疫吸附试验(double-antibody sandwich enzyme linked immunosorbent assay, DAS-ELISA)，用于快速定量检测SCRV灭活疫苗的抗原浓度。DAS-ELISA能否用于SCRV的临床样品检测还有待进一步研究。PCR和荧光定量PCR作为传统的病毒分子生物学检测方法具有重要应用价值，但这2种方法都需要价格昂贵的热循环仪和专业人员操作，通常要在实验室里才能完成检测。一些不需要热循环，更适合于现场检测的等温扩增技术，如环介导等温扩增、指数扩增反应和催化发夹自组装等，以及基于免疫学方法的胶体金检测试纸条等技术在SCRV检测方面的应用研究尚未见报道。值得指出的是，上述已建立的各种检测方法在特异性方面仅能区分不同病毒属病原体，对于能够有效区分同病毒属内的病毒(如MSRV、HSHRV、SHVV和CrERV)的检测方法尚未见报道。特异性达到能区分病毒种之间差异的检测方法不仅对于SCRV临床检测具有重要价值，对于流行病毒调查也有重要意义。建立特异性更高的SCRV检测方法(如针对特定抗原位点的单克隆抗体法)亟待解决。

5 SCRV防控策略

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5.1 SCRV抗病毒药物

药物和疫苗是治疗和控制病原传播的重要手段。有关SCRV的抗病毒药物研究主要有源于组织因子途径抑制剂的多肽^[72]、三联基序家族蛋白中的鳜鱼三联基序59^[73]、巨胞饮化学阻断剂乙基异丙基阿米洛利^[74]、天然二氢黄酮类化合物橙皮素^[75]和真核翻译延伸因子1α^[76]。上述几类药物在实验室条件下对SCRV的感染有抑制效应。此外，一些基于感染后的细胞信号通路研究也为SCRV药物研究提供了潜在的抗病毒靶点^[77-79]。最近，Yu等^[80]利用基因编辑技术CRISPR/Cas13d系统，通过设计多组靶向SCRV基因组或其mRNA的向导RNA (guide RNA, gRNA)，筛选出5组gRNA可在细胞水平上有效抑制SCRV的感染，这为研发防控鱼类弹状病毒的对策提供了新思路。对于鱼类病毒药物的研发，与大型哺乳动物和人类抗病毒药物研发有一个显著区别，即药物的成本问题。鱼群一旦发病，药物治疗成本和病害经济损失之间的平衡是重要考量因素，而当前研究尚未将这一重要影响因素纳入考量。

5.2 SCRV疫苗

对于水生动物而言，传染病的预防价值重于治疗。疫苗是传染性疾病预防的关键要素。疫苗的类别主要有灭活疫苗、弱毒疫苗、重组蛋白疫苗、DNA疫苗和近年新兴的mRNA疫苗等。至今，尚无可用于鳜鱼养殖的SCRV疫苗，有关SCRV的疫苗研究也较少。已报道的SCRV疫苗研究仅有DNA疫苗、灭活疫苗和弱毒疫苗。DNA疫苗是将SCRV的G基因构建到真核表达载体pCDNA3.1中，通过肌内注射进行免疫，基于G基因的DNA疫苗对鳜鱼的死亡保护率为77.5%^[81]。灭活疫苗是将SCRV强毒株SCRV-GM1503株用甲醛灭活，制备的灭活疫苗通过腹腔注射途径免疫，对鳜鱼的死亡保护率为84%^[82]。弱毒疫苗是分离自大口黑鲈的一株弱毒株MSRV-SS，通过腹腔注射进行免疫后，对SCRV强毒株SCRV-GM攻毒有很好的交叉保护作用，死亡保护率为100%^[5,83]。然而，MSRV-SS株作为弱毒活疫苗，通过浸泡免疫方式进行免疫的保护效果尚不清楚。

如前所述的各类疫苗，除弱毒疫苗外，其他类别的疫苗都需采用注射的方式进行免疫。注射方式免疫费时费力。与其他类型疫苗相比，弱毒疫苗不仅具有良好的免疫原性，而且可以采用浸泡的方式进行免疫。因此，对于鱼类而言，弱毒疫苗具有独特的优势。然而，获得弱毒毒株的途径多局限于将致病的强毒株在体外进行数十次或更多的传代以致弱病毒，或从自然界分离获得天然的弱毒株。这种获取弱毒株的方法不仅耗时费力，而且具有非常大的偶然性，无法快速应对传染病的发生。近期建立的SCRV反向遗传系统为人工快速致弱SCRV毒株提供了技术平台，将极大地促进SCRV疫苗的研究和开发。

5.3 环境因素对SCRV防控的影响

传染病的发生和流行是“病原-环境-宿主”三者相互作用的结果。对于水生动物病毒而言，环境因素是驱动很多鱼类病毒病发生与发展的关键要素。在各种环境要素中，温度扮演着核心作用。一方面，水生动物病毒复制和致病性都具有严格的温度依赖性；另一方面，病毒在环境中的稳定性和存活时间受温度的影响极大，而病毒的稳定性和存活时间又是影响病毒传播的重要中间环节。明确病毒在水体和固体介质表面等各种环境下的存活时间对病毒的消杀和病原的有效控制具有重要指导意义。最近的研究发现，SCRV在液体环境中时，25 ℃条件下可存活长达近半年，4 ℃的环境下表现出更为明显的稳定性，病毒滴度在一年内都维持在较高水平；SCRV在玻璃、不锈钢和塑料介质表面均能存活较长时间；采用常规的55 ℃、30 min热灭活病毒方法不适用于SCRV，需要60 ℃处理30 min才能被完全失活^[84]。这要比哺乳动物弹状病毒水泡性口炎病毒在同样温度下稳定得多，水泡性口炎病毒在55 ℃条件下4 min即可完全灭活^[85]。除上述开展的SCRV稳定性研究外，其他环境要素对SCRV的防控影响还知之甚少，有待进一步研究。

6 SCRV目前研究问题和未来研究展望

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6.1 SCRV流行病学调查亟待解决

鳜鱼是我国传统的养殖品种，有上千年的养殖历史。我国鳜鱼养殖主要分布在广东、湖北和安徽三省，分别占全国总产量的31.48%、23.64%和12.74%；鳜鱼苗种集中在广东省，占全国95%以上^[1]。GenBank数据库提交的SCRV基因序列来源地除由浙江省分离的S3株外，其他毒株均来自广东省。流行病学调查包括血清学调查和病原学调查，通过流行病学调查不仅对于揭示病毒变异情况、为研发高保护率疫苗奠定基础具有重要意义，而且对于追溯疫病来源、明确病原分布、阐明病毒传播动力学、评估疫情风险、实施精准隔离和制定疫病管理措施也具有重要意义。此外，SCRV也能感染大口黑鲈，分离自大口黑鲈的弹状病毒称之为MSRV。虽然SCRV在文献报道中的出现比MSRV更早一些，但SCRV和MSRV在进化关系上哪种病毒出现得更早以及二者之间的进化关系依旧是个未知问题。大口黑鲈是由北美引进的品种，最早于1983年引入我国。截至2025年9月，所有有关SCRV的研究文献均来自我国，其在国外的感染和分布情况，尤其是大口黑鲈原产地北美的感染情况尚不清楚。广泛而深入的SCRV血清学和病原学调查对于上述问题的阐明具有重要价值。然而，至今尚无针对SCRV流行病学的系统调查文献，其流行病学调查亟待解决。

6.2 SCRV的致病机制有待揭示

揭示病毒的致病机制对于开发抗病毒药物和提出病毒预防策略具有重要意义。然而，鲜有关SCRV致病机制的报道，其致病机制还有待阐明。例如，SCRV的受体是什么，病毒进入体内的主要途径是什么，病毒如何进入靶细胞和靶器官、其病理进程如何，造成组织器官损伤和功能障碍的机理是什么，不同毒株毒力差异的原因是什么，病毒的毒力决定基因是什么等基础性问题都有待揭示。SCRV对不同年龄段的鳜鱼致病情况如何，对其他鱼种是否致病等也是待解决的关键问题。此外，除了病毒和宿主自身情况外，鱼类疾病暴发的重要诱因和病情发展受环境因素的影响非常大，如温度、水质和日照等外部因素对SCRV病情的影响如何以及影响机制是什么，多角度多变量因子系统性研究SCRV致病机制还有待开展。

6.3 SCRV的基础研究和应用研究需要平衡

病毒学研究包括基础研究和应用研究两大方面。SCRV的研究主要集中在基础研究，国家和各省级单位支持的相关基金项目也大都集中在基础研究，这导致有关SCRV的应用研究相对较少。对于鱼类病毒而言，发病后的治疗价值不大，更应注重应用疫苗进行防控。当前有关SCRV的疫苗研究还非常薄弱。未来对SCRV疫苗的相关应用研究，除了利用现代生物技术工具开发具有良好免疫原性和免疫反应性的疫苗类别外，疫苗佐剂和疫苗递送系统研究也具有重要意义。由于鱼类非特异性免疫在自身免疫反应中较哺乳动物起着更为重要的作用，利用对非特异性免疫反应有较强刺激作用的佐剂可能会起到更好的保护效果。此外，由于鱼类生长环境的特殊性，开发能够通过浸泡或口服途径进行免疫的疫苗递送系统也具有重要价值。上述这些具有实用价值和现实意义的应用研究还非常少，未来有待向这方面平衡和加强。

整体而言，水生动物病毒学的研究落后于哺乳动物和人类病毒学，SCRV的研究也是如此。有关SCRV的研究大都借用哺乳动物或人类病毒学的研究思路或方法，将相关内容延伸到SCRV的研究，而基于SCRV本身特征或应用SCRV作为模型进行的原创性研究较少。SCRV自发现至今已近30年历史，在一些方面取得了进步，尤其是现代测序技术的应用极大地推动了SCRV与宿主抗病毒反应之间关系的研究。SCRV反向遗传系统的建立也将有助于推动SCRV弱毒疫苗和SCRV基因组功能研究。然而，有关SCRV的流行病学和致病机制研究还非常欠缺，有待于进一步加强相关研究。

基金

收起

浙江省自然科学基金(LY14C180001)
绍兴文理学院人才引进项目(13011001002/245)

参考文献

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文献

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参考文献引证文献

排序方式：

[1]

国家特色淡水鱼产业技术体系. 中国鳜鱼产业发展报告[J]. 中国水产, 2021(4): 23-32.

[2]

ZHANG

, KE

, GUI

, ZHAO

. Recent insights into aquatic viruses: emerging and reemerging pathogens, molecular features, biological effects, and novel investigative approaches[J]. Water Biology and Security, 2022, 1(4): 100062.

[3]

张奇亚, 李正秋. 在患病鳜鱼组织中观察到3种病毒[J]. 科学通报, 1999，44(2): 192-195.

[4]

, LIN

, LIANG

, LIU

, HUANG

, LI

, SU

. The biological features and genetic diversity of novel fish rhabdovirus isolates in China[J]. Archives of Virology, 2017, 162(9): 2829-2834.

[5]

张莉娟. 鳜弹状病毒病减毒活疫苗候选株的筛选、纯化及其免疫原性分析[D]. 上海: 上海海洋大学, 2018.

ZHANG

. Screening, purification and immunogenicity analysis of a candidate of live-attenuated vaccine for Siniperca chuatsi rhabdovirus disease[D]. Shanghai: Shanghai Ocean University, 2018 (in Chinese).

[6]

, DENG

, BAI

, LI

, YU

, QUAN

, YANG

, JIANG

, ZHU

, YE

. A strain of Siniperca chuatsi rhabdovirus causes high mortality among cultured largemouth bass in South China[J]. Journal of Aquatic Animal Health, 2013, 25(3): 197-204.

[7]

TAO

, GUI

, ZHANG

. Isolation and characterization of a rhabdovirus from co-infection of two viruses in mandarin fish[J]. Aquaculture, 2007, 262(1): 1-9.

[8]

TAO

, ZHOU

, GUI

, ZHANG

. Genomic sequence of mandarin fish rhabdovirus with an unusual small non-transcriptional ORF[J]. Virus Research, 2008, 132(1/2): 86-96.

[9]

陶建军. 鳜鱼弹状病毒的分离、鉴定及基因组测序分析[D]. 武汉: 中国科学院研究生院(水生生物研究所), 2006.

TAO

. Isolation, identification and complete genomic sequence analysis of the mandarin fish Siniperca chuatsi rhabdovirus[D]. Wuhan: Institute of Hydrobiology, Chinese Academy of Sciences, 2006 (in Chinese).

[10]

KUHN

, ADKINS

, ALKHOVSKY

, AN

, AVŠIČ-ŽUPANC

, AYLLÓN

, BAČNIK

, BAHL

, BALKEMA-BUSCHMANN

, BALLINGER

, BEER

, BEJERMAN

, BERGERON

, BIEDENKOPF

, BLAIR

, BLASDELL

, BRADFUTE

, BRIESE

, BROWN

, et al. Annual (2024) taxonomic update of RNA-directed RNA polymerase-encoding negative-sense RNA viruses (realm Riboviria: Kingdom Orthornavirae: Phylum Negarnaviricota)[J]. Journal of General Virology, 2025, 106(6): 002077.

[11]

LYU SJ, YUAN

, ZHANG

, SHI

, HANG

, LIU

, WU

. Isolation and characterization of a novel strain (YH01) of Micropterus salmoides rhabdovirus and expression of its glycoprotein by the baculovirus expression system[J]. Journal of Zhejiang University Science B, 2019, 20(9): 728-739.

[12]

QIN

, ZHANG

, GUO

, DENG

, LIU

, YAO

. Isolation and identification of a new strain Micropterus salmoides rhabdovirus (MSRV) from largemouth bass Micropterus salmoides in China[J]. Aquaculture, 2023, 572: 739538.

[13]

, YI

, CHENG

, YANG

, FEI

. A concise review on advancement of Micropterus salmoides rhabdovirus (MSRV): current status and challenges[J]. Aquaculture Reports, 2023, 30: 101551.

[14]

ZENG

, WANG

, LIU

, LIANG

, FANG

, WU

. Genomic characterization and taxonomic position of a rhabdovirus from a hybrid snakehead[J]. Archives of Virology, 2014, 159(9): 2469-2473.

[15]

DENG

, ZHENG

, LI

, MO

, ZHAO

, LUO

, YIN

, SHI

, WANG

. Isolation and identification of hybrid snakehead rhabdovirus (HSHRV) and its immune response in the hybrid Snakehead (male Channa argus×female Channa maculata)[J]. Microbial Pathogenesis, 2024, 196: 106983.

[16]

LIU

, WEN

, HU

, WANG

, LIANG

, LI

, VAKHARIA

, LIN

. Breaking the host range: mandarin fish is susceptible to a vesiculovirus derived from snakehead fish[J]. Journal of General Virology, 2015, 96(Pt 4): 775-781.

[17]

OU T, ZHU

, CHEN

, ZHANG

. Isolation and identification of a lethal rhabdovirus from farmed rice field eels Monopterus albus [J]. Diseases of Aquatic Organisms, 2013, 106(3): 197-206.

[18]

LIU

, FAN

, LI

, ZHAO

, ZHOU

, JIANG

, ZENG

, CAIN

, ZENG

. Isolation, identification, and classification of a novel rhabdovirus from diseased Chinese rice-field eels (Monopterus albus)[J]. Archives of virology, 2019, 164(1): 105-116.

[19]

CHEN

, HUO

, JIN

, PENG

, LI

, WU

, ZHANG

, TIAN

, WANG

, LI

, LIU

, NIE

. The infectious haemorrhagic syndrome virus (IHSV) from rice-field eel (Monopterus albus): isolation, genome sequence, cross-infection and induced-immune response in Chinese perch (Siniperca chuatsi)[J]. Aquaculture, 2024, 583: 740561.

[20]

WALKER

, BIGARRÉ

, KURATH

, DACHEUX

, PALLANDRE

. Revised taxonomy of rhabdoviruses infecting fish and marine mammals[J]. Animals, 2022, 12(11): 1363.

[21]

RAMASWAMY

, RASHID

, RAMASAMY

, JAYAVELU

, VENKATARAMAN

. Revisiting viral RNA-dependent RNA polymerases: insights from recent structural studies[J]. Viruses, 2022, 14(10): 2200.

[22]

BIACCHESI

, THOULOUZE

, BÉARZOTTI

, YU

, BRÉMONT

. Recovery of NV knockout infectious hematopoietic necrosis virus expressing foreign genes[J]. Journal of Virology, 2000, 74(23): 11247-11253.

[23]

AMMAYAPPAN

, KURATH

, THOMPSON

, VAKHARIA

. A reverse genetics system for the great lakes strain of viral hemorrhagic septicemia virus: the NV gene is required for pathogenicity[J]. Marine Biotechnology, 2011, 13(4): 672-683.

[24]

JOHNSON

, SIMON

, KIM

, LEONG

. Production of recombinant snakehead rhabdovirus: the NV protein is not required for viral replication[J]. Journal of Virology, 2000, 74(5): 2343-2350.

[25]

RYU SJ, KWAK

, KIM

. Effect of NV gene deletion in the genome of hirame rhabdovirus (HIRRV) on viral replication and the type I interferon response of the host cell[J]. Archives of Virology, 2022, 167(1): 77-84.

[26]

SOUTO

, LAMA

, MÉROUR

, MEHRAZ

, BERNARD

, LAMOUREUX

, MASSAAD

, FRÉTAUD

, RIGAUDEAU

, MILLET

, LANGEVIN

, BIACCHESI

. In vivo multiscale analyses of spring viremia of carp virus (SVCV) infection: from model organism to target species[J]. PLoS Pathogens, 2024, 20(8): e1012328.

[27]

FENG

, SU

, LIN

, TU

. Development of a reverse genetics system for snakehead vesiculovirus (SHVV)[J]. Virology, 2019, 526: 32-37.

[28]

LIU

, ZHANG

, XU

, HUANG

, ZHONG

, ZHU

, WEI

. Recovery of Siniperca chuatsi rhabdovirus from cloned cDNA[J]. Journal of Fish Diseases, 2023, 46(2): 127-135.

[29]

LIU

, HUANG

, MIAO

, LI

, WANG

, XU

, ZHANG

, WEI

. Infection tracing and organ tropism of Siniperca chuatsi rhabdovirus expressing enhanced green fluorescent protein[J]. Aquaculture, 2023, 574: 739684.

[30]

LIU

, ZHANG

, XU

, HUANG

, ZHONG

, ZHU

, WEI

. Isolation, genomic and biological characterizations of a rhabdovirus from mandarin fish (Siniperca chuatsi)[J]. Aquaculture, 2023, 563: 738894.

[31]

DONG

, XIE

, LI

, XU

, SUN

. RNF34 negatively regulates innate immunity via ubiquitin-mediated TAK1 degradation in miiuy croaker (Miichthys miiuy)[J]. Fish & Shellfish Immunology, 2025, 166: 110578.

[32]

LUO

, FU

, ZHANG

, LIANG

, NIU

, LIN

, MA

, LIU

, LI

. Development of a new marine fish continuous cell line derived from brain of red sea bream (Pagrosomus major) and its application to fish virology and heavy metal toxicology[J]. Animals, 2023, 13(22): 3524.

[33]

ZHANG

, SUN

, TAN

, GONG

, KONG

, LIU

. Transcriptomic profiling of the immune response to Siniperca chuatsi rhabdovirus (SCRV) in E11 cells[J]. Aquaculture International, 2024, 32(6): 7821-7834.

[34]

HUANG

, WANG

, GAO

, WU

, ZHANG

, SHI

, DING

, ZHAO

. Transcriptome analysis of immune response against Siniperca chuatsi rhabdovirus infection in mandarin fish Siniperca chuatsi [J]. Journal of Fish Diseases, 2021, 44(6): 675-687.

[35]

, MENG

, ZHANG

. Siniperca chuatsi rhabdovirus (SCRV)-induced key pathways and major antiviral genes in fish cells[J]. Microorganisms, 2022, 10(12): 2464.

[36]

SIOUNING

, SEONG

, KONDO

, BHASSU

. microRNA regulation in infectious diseases and its potential as a biosensor in future aquaculture industry: a review[J]. Molecules, 2023, 28(11): 4357.

[37]

ZHANG

, TU

, ZHANG

. microRNA regulation of viral replication in teleost fish: a review[J]. Reviews in Aquaculture, 2021, 13(3): 1367-1378.

[38]

ZHANG

, CHU

, CHANG

, XU

. Inducible microRNA-217 inhibits NF-κB-and IRF3-driven immune responses in lower vertebrates through targeting TAK1[J]. Journal of Immunology, 2020, 205(6): 1620-1632.

[39]

ZHENG

, CHU

, XU

. The long noncoding RNA NARL regulates immune responses via microRNA-mediated NOD1 downregulation in teleost fish[J]. Journal of Biological Chemistry, 2021, 296: 100414.

[40]

, CHU

, CUI

, BI

. Inducible microRNA-3570 feedback inhibits the RIG-I-dependent innate immune response to rhabdovirus in teleost fish by targeting MAVS/IPS-1[J]. Journal of Virology, 2018, 92(2): e01594-17.

[41]

CHU

, XU

, ZHENG

, CHANG

, ZHANG

. Long noncoding RNA MARL regulates antiviral responses through suppression miR-122-dependent MAVS downregulation in lower vertebrates[J]. PLoS Pathogens, 2020, 16(7): e1008670.

[42]

GAO

, CHANG

, SUN

, XU

. microRNA-2187 modulates the NF-κB and IRF3 pathway in teleost fish by targeting TRAF6[J]. Frontiers in Immunology, 2021, 12: 647202.

[43]

CHANG

, ZHENG

, SUN

, GENG

, XU

. Long noncoding RNA MIR2187HG suppresses TBK1-mediated antiviral signaling by deriving miR-2187-3p in teleost fish[J]. Journal of Virology, 2022, 96(1): e0148421.

[44]

SUN

, ZHANG

, HONG

, ZHENG

, CUI

, LIU

, XU

. microRNA-181b-2 and microRNA-21-1 negatively regulate NF-κB and IRF3-mediated innate immune responses via targeting TRIF in teleost[J]. Frontiers in Immunology, 2021, 12: 734520.

[45]

, CHU

, CUI

. Rhabdovirus-inducible microRNA-210 modulates antiviral innate immune response via targeting STING/MITA in fish[J]. Journal of Immunology, 2018, 201(3): 982-994.

[46]

ZHAO

, LIN

, LI

, BABU

, FU

, LIU

, LIANG

, LIU

, LIN

. microRNAs profiles of Chinese Perch Brain (CPB) cells infected with Siniperca chuatsi rhabdovirus (SCRV)[J]. Fish & Shellfish Immunology, 2019, 84: 1075-1082.

[47]

ZHENG

, CHANG

, LUO

, LIU

, XU

. The long noncoding RNA MIR122HG is a precursor for miR-122-5p and negatively regulates the TAK1-induced innate immune response in teleost fish[J]. Journal of Biological Chemistry, 2022, 298(4): 101773.

[48]

CUI

, ZHENG

, XU

, SUN

. Long noncoding RNA MIR122HG inhibits MAVS-mediated antiviral immune response by deriving miR-122 in miiuy croaker (Miichthys miiuy)[J]. Viruses, 2022, 14(5): 930.

[49]

CHU

, XU

, ZHENG

, CHANG

, ZHANG

. Long noncoding RNA AANCR modulates innate antiviral responses by blocking miR-210-dependent MITA downregulation in teleost fish, Miichthys miiuy [J]. Science China Life Sciences, 2021, 64(7): 1131-1148.

[50]

CHU

, ZHENG

, SU

, ZHANG

, CHANG

, GAO

, XU

. A highly conserved circular RNA circRasGEF1B enhances antiviral immunity by regulating miR-21-3p/MITA pathway in lower vertebrates[J]. Journal of Virology, 2021, 95(7): e02145-20.

[51]

, ZHENG

, GENG

, CUI

, TAO

, XU

. circCBL and its host gene CBL collaboratively enhance the antiviral immunity and antibacterial immunity by targeting MITA in fish[J]. Journal of Virology, 2023, 97(10): e0104623.

[52]

ZHENG

, CHU

, YANG

, SUN

, XU

. Circular RNA circDtx1 regulates IRF3-mediated antiviral immune responses through suppression of miR-15a-5p-dependent TRIF downregulation in teleost fish[J]. PLoS Pathogens, 2021, 17(3): e1009438.

[53]

, ZHENG

, PAN

, LV

, XIN

, XU

. Circular RNA circSamd4a regulates antiviral immunity in teleost fish by upregulating STING through sponging miR-29a-3p[J]. Journal of Immunology, 2021, 207(11): 2770-2784.

[54]

ZHENG

, SU

, LV

, XIN

, XU

. Exon-intron circular RNA circRNF217 promotes innate immunity and antibacterial activity in teleost fish by reducing miR-130-3p function[J]. Journal of Immunology, 2022, 208(5): 1099-1114.

[55]

ZHENG

, SUN

, YANG

, XU

. The circular RNA circBCL2L1 regulates innate immune responses via microRNA-mediated downregulation of TRAF6 in teleost fish[J]. Journal of Biological Chemistry, 2021, 297(4): 101199.

[56]

, CHU

, ZHENG

, CHANG

, GAO

, ZHANG

, XU

. Circular RNA circPIKfyve acts as a sponge of miR-21-3p to enhance antiviral immunity through regulating MAVS in teleost fish[J]. Journal of Virology, 2021, 95(8): e02296-20.

[57]

WANG

, SONG

, YAN

, XU

. A novel protein encoded by circVPS13D attenuates antiviral innate immunity by targeting MAVS in teleost fish[J]. Journal of Virology, 2023, 97(11): e0088623.

[58]

WANG

, ZHENG

, LV

, SONG

, XU

. circMORC3-encoded novel protein negatively regulates antiviral immunity through synergizing with host gene MORC3 [J]. PLoS Pathogens, 2023, 19(12): e1011894.

[59]

ZHENG

, WANG

, GENG

, XU

. CircYthdc2 generates polypeptides through two translation strategies to facilitate virus escape[J]. Cellular and Molecular Life Sciences, 2024, 81(1): 91.

[60]

ZHENG

, ZHU

, LUO

, ZHAO

, XU

. A novel protein NLRP12-119aa that prevents rhabdovirus replication by disrupting the RNP complex formation[J]. Advanced Science, 2025, 12(12): e2409953.

[61]

, HE

, YANG

, LI

, WANG

. Unveiling the hidden regulators: the impact of lncRNAs on zoonoses[J]. International Journal of Molecular Sciences, 2024, 25(6): 3539.

[62]

VIERBUCHEN

, FITZGERALD

. Long non-coding RNAs in antiviral immunity[J]. Seminars in Cell & Developmental Biology, 2021, 111: 126-134.

[63]

WEI

, FAN

, MAO

. A review on circular RNA translation and its implications in disease[J]. Methods in Molecular Biology, 2025, 2883: 109-137.

[64]

HUSSEN

, ABDULLAH

, KHUDHUR

, SAMSAMI

, TAHERI

. Circular RNAs as a therapeutic modality for viral infections and innovative strategies to overcome key challenges: a review[J]. International Journal of Biological Macromolecules, 2025, 318: 145088.

[65]

YIN

, LI

, ZHOU

. Circular RNAs in viral infection and antiviral treatment[J]. Cells, 2024, 13(23): 2033.

[66]

梁红茹, 范芷仪, 蔡秀珠, 付小哲, 林强, 刘礼辉, 黄志斌, 牛银杰, 林蠡, 李宁求. 鳜弹状病毒与传染性脾肾坏死病毒双重PCR检测方法的建立[J]. 西北农林科技大学学报(自然科学版), 2020, 48(3): 39-46, 53.

LIANG

, FAN

, CAI

, FU

, LIN

, LIU

, HUANG

, NIU

, LIN

, LI

. Establishment of duplex PCR assay for detecting Siniperca chuatsi rhabdovirus and infectious spleen and kidney necrosis virus[J]. Journal of Northwest A&F University (Natural Science Edition), 2020, 48(3): 39-46, 53 (in Chinese).

[67]

梁红茹, 黄瑜聪, 付小哲, 林强, 刘礼辉, 牛银杰, 黄志斌, 林蠡, 李宁求. 鳜蛙病毒及鳜弹状病毒双重PCR检测方法的建立及应用[J]. 中国预防兽医学报, 2021, 43(3): 290-295.

LIANG

, HUANG

, FU

, LIN

, LIU

, NIU

, HUANG

, LIN

, LI

. Development of a duplex PCR for the detection of Siniperca chuatsi ranairidovirus and Siniperca chuatsi rhabdovirus[J]. Chinese Journal of Preventive Veterinary Medicine, 2021, 43(3): 290-295 (in Chinese).

[68]

梁红茹, 马赛亚, 付小哲, 林强, 刘礼辉, 牛银杰, 黄志斌, 林蠡, 李宁求. 传染性脾肾坏死病毒、鳜鱼蛙病毒和鳜弹状病毒三重PCR检测方法的建立[J]. 西北农林科技大学学报(自然科学版), 2021, 49(11): 17-24.

LIANG

, MA

, FU

, LIN

, LIU

, NIU

, HUANG

, LIN

, LI

. Establishment of multiple PCR assay for detecting infectious spleen and kidney necrosis virus, Siniperca chuatsi ranairidovirus and Siniperca chuatsi rhabdovirus[J]. Journal of Northwest A&F University (Natural Science Edition), 2021, 49(11): 17-24 (in Chinese).

[69]

梁红茹, 蔡秀珠, 范芷仪, 林强, 付小哲, 刘礼辉, 黄志斌, 牛银杰, 林蠡, 李宁求. 鳜弹状病毒TaqMan荧光定量PCR检测方法的建立及应用[J]. 中国预防兽医学报, 2019, 41(9): 929-934.

LIANG

, CAI

, FAN

, LIN

, FU

, LIU

, HUANG

, NIU

, LIN

, LI

. Establishment and application of a TaqMan real-time PCR assay for the detection of Siniperca chuatsi rhabdovirus[J]. Chinese Journal of Preventive Veterinary Medicine, 2019, 41(9): 929-934 (in Chinese).

[70]

LIN

, ZHAO

, FU

, LIU

, LIANG

, NIU

, CHEN

, HUANG

, LIN

, LI

. Development of strand-specific real-time RT-PCR for the analysis of SCRV transcription and replication dynamics[J]. Microbial Pathogenesis, 2019, 129: 146-151.

[71]

NIU

, ZHANG

, WANG

, LI

, LIN

, LIU

, LIANG

, HUANG

, FU

. Development of double-antibody sandwich ELISA for rapidly quantitative detection of antigen concentration in inactivated SCRV vaccine[J]. Aquaculture, 2020, 520: 734671.

[72]

, LI

, ZHANG

, FU

, LIANG

, NIU

, LIN

, LUO

, LIU

, SU

, ZHOU

, LI

. Tissue factor pathway inhibitors disrupt structures of rhabdovirus/ranairidovirus and inhibit viral infection in Chinese perch, Siniperca chuatsi [J]. Fish & Shellfish Immunology, 2024, 149: 109553.

[73]

NIU

, FU

, LIN

, LIANG

, LUO

, ZUO

, LIU

, LI

. The composition and antiviral activity of scTRIM59 in Mandarin fish[J]. Fish & Shellfish Immunology, 2022, 130: 86-92.

[74]

LIANG

, LUO

, LIN

, FU

, NIU

, MA

, LI

. EIPA, A macropinocytosis blocking agent, inhibits Siniperca chuatsi rhabdovirus infection in vitro and in vivo [J]. Microbial Pathogenesis, 2025, 206: 107751.

[75]

, SUN

, MA

, ZHOU

. Hesperetin inhibits Siniperca chuatsi rhabdovirus replication by reducing mitochondrial apoptosis in EPC cells[J]. Israeli Journal of Aquaculture - Bamidgeh, 2025, 77(1): 213-222.

[76]

MENG

, JIANG

, YU

, ZHANG

, KE

. Eukaryotic translation elongation factor 1 alpha (eEF1A) inhibits Siniperca chuatsi rhabdovirus (SCRV) infection through two distinct mechanisms[J]. Journal of Virology, 2023, 97(11): e0122623.

[77]

, FU

, LUO

, LIN

, LIANG

, NIU

, LIU

, LI

. Role of asparagine biosynthesis pathway in Siniperca chuatsi rhabdovirus proliferation[J]. Frontiers in Microbiology, 2023, 14: 1165491.

[78]

NIU

, YANG

, LIANG

, LUO

, MA

, LIN

, FU

, LI

. scTRIM44 positively regulated Siniperca chuatsi rhabdovirus through RIG-I- and MDA5-mediated interferon signaling[J]. Viruses, 2024, 16(12): 1876.

[79]

, CHEN

, WANG

, LI

, PANG

, LIU

, LI

, HOU

, WANG

, NIE

. Interaction of nmi and IFP35 promotes mutual protein stabilization and IRF3 and IRF7 degradation to suppress type I IFN production in teleost fish[J]. The Journal of Immunology, 2023, 210(10): 1494-1507.

[80]

, ZHANG

, KE

. CRISPR/Cas13d mediated inhibition of Siniperca chuatsi rhabdovirus infection in fish cells[J]. Aquaculture, 2025, 602: 742355.

[81]

CHEN

, LEI

, ZHANG

. The antiviral defense mechanisms in mandarin fish induced by DNA vaccination against a rhabdovirus[J]. Veterinary Microbiology, 2012, 157(3/4): 264-275.

[82]

罗霞, 付小哲, 林强, 刘礼辉, 牛银杰, 梁红茹, 李宁求. 鳜鱼传染性脾肾坏死病和弹状病毒病二联灭活疫苗毒种及种子批的研究[J]. 西北农林科技大学学报(自然科学版), 2022, 50(1): 1-9.

LUO

, FU

, LIN

, LIU

, NIU

, LIANG

, LI

. Virus seed and seed batches of mandarin fish ISKNV and SCRV bivalent inactivated vaccine[J]. Journal of Northwest A&F University (Natural Science Edition), 2022, 50(1): 1-9 (in Chinese).

[83]

ZHANG

, LI

, LIN

, LIU

, LIANG

, HUANG

, FU

. An avirulent Micropterus salmoides rhabdovirus vaccine candidate protects Chinese perch against rhabdovirus infection[J]. Fish & Shellfish Immunology, 2018, 77: 474-480.

[84]

, HUANG

, ZHONG

, ZHU

, LIU

, WEI

. Thermal and environmental stability of Siniperca chuatsi rhabdovirus[J]. Aquaculture, 2023, 568: 739308.

[85]

ZIMMER

, SUMMERMATTER

, ZIMMER

. Stability and inactivation of vesicular stomatitis virus, a prototype rhabdovirus[J]. Veterinary Microbiology, 2013, 162(1): 78-84.

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doi: 10.13343/j.cnki.wsxb.20250576

接收时间：2025-07-24
首发时间：2026-01-12
出版时间：2026-01-04

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收稿日期：2025-07-24
录用日期：2025-09-23

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Natural Science Foundation of Zhejiang Province(LY14C180001)

浙江省自然科学基金(LY14C180001)

Shaoxing University Talent Introduction Program(13011001002/245)

绍兴文理学院人才引进项目(13011001002/245)

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绍兴文理学院医学院，浙江绍兴

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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TxT

Virus name	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18
1	100	96.8	97.4	97.4	92.7	97.3	97.5	97.6	97.1	96.9	97.3	93.1	93.2	91.9	92.1	93.1	92.9	92.3
2		100	96.1	96.1	94.6	96.1	96.2	96.4	95.9	95.5	96.0	94.9	95.0	93.4	93.8	95.0	94.6	94.1
3			100	99.3	92.3	97.9	99.3	98.3	99.1	98.8	99.2	92.7	92.7	91.5	91.8	92.6	92.4	92.0
4				100	92.3	98.0	99.2	98.4	99.5	99.5	99.6	92.7	92.8	91.4	91.7	92.6	92.4	92.1
5					100	92.1	92.2	92.3	91.9	91.7	92.2	93.3	93.3	92.0	92.4	93.1	92.9	92.5
6						100	98.1	99.4	97.8	97.5	98.2	92.6	92.7	91.4	91.7	92.5	92.3	91.8
7							100	98.6	99.1	98.8	99.2	92.8	92.8	91.5	91.8	92.6	92.4	92.1
8								100	98.2	98.0	98.3	92.9	93.0	91.7	92.0	92.8	92.5	92.1
9									100	99.0	99.5	92.5	92.5	91.2	91.5	92.3	92.1	91.8
10										100	99.2	92.2	92.3	91.0	91.3	92.1	91.9	91.6
11											100	92.7	92.7	91.4	91.7	92.6	92.4	92.0
12												100	99.9	96.9	97.3	93.5	93.2	92.8
13													100	96.9	97.3	93.5	93.2	92.8
14														100	98.9	92.4	92.0	91.6
15															100	92.7	92.3	91.9
16																100	98.2	94.5
17																	100	94.1
18																		100

Virus name

SCRV

MSRV

HSRV

SHVV

CrERV

IHSV

100

96.8

97.4

92.7

97.3

97.5

97.6

97.1

96.9

97.3

93.1

93.2

91.9

92.1

93.1

92.9

92.3

100

96.1

94.6

96.1

96.2

96.4

95.9

95.5

96.0

94.9

95.0

93.4

93.8

95.0

94.6

94.1

100

99.3

92.3

97.9

99.3

98.3

99.1

98.8

99.2

92.7

91.5

91.8

92.6

92.4

92.0

100

92.3

98.0

99.2

98.4

99.5

99.6

92.7

92.8

91.4

91.7

92.6

92.4

92.1

100

92.1

92.2

92.3

91.9

91.7

92.2

93.3

92.0

92.4

93.1

92.9

92.5

100

98.1

99.4

97.8

97.5

98.2

92.6

92.7

91.4

91.7

92.5

92.3

91.8

100

98.6

99.1

98.8

99.2

92.8

91.5

91.8

92.6

92.4

92.1

100

98.2

98.0

98.3

92.9

93.0

91.7

92.0

92.8

92.5

92.1

100

99.0

99.5

92.5

91.2

91.5

92.3

92.1

91.8

100

99.2

92.2

92.3

91.0

91.3

92.1

91.9

91.6

100

92.7

91.4

91.7

92.6

92.4

92.0

100

99.9

96.9

97.3

93.5

93.2

92.8

100

96.9

97.3

93.5

93.2

92.8

100

98.9

92.4

92.0

91.6

100

92.7

92.3

91.9

100

98.2

94.5

100

94.1

100

RNA type	RNA name	Targeting molecule	Function mode	References
miRNA	miR-217	TGF-b-activated kinase 1 (TAK1)	Inhibition of protein expression (IPE)	[38]
	miR-217-5p	Nucleotide oligomerization domain 1 (NOD1)	IPE	[39]
	miR-3570	Mitochondrial antiviral signaling protein (MAVS)	IPE	[40]
	miR-122	MAVS	IPE	[41]
	miR-2187	Tumor necrosis factor receptor related factor 6 (TRAF6)	IPE	[42]
	miR-2187-3p	TANK-binding kinase 1 (TBK1)	IPE	[43]
	miR-181b-2	TIR-domain-containing adapter-inducing interferon-b (TRIF)	IPE	[44]
	miR-21-1	TRIF	IPE	[44]
	miR-210	Stimulator of IFN genes (STING)	IPE	[45]
	miR-214	N and P genes of SCRV	Inhibition of viral replication	[46]
lncRNA	MIR2187HG	miR-2187-3p	Pre-miRNA	[43]
	MIR122HG	miR-122, miR-122-5p	Pre-miRNA	[47-48]
	lncRNA AANCR	miR-210	Sponge effect	[49]
	lncRNA MARL	miR-122	Sponge effect	[41]
	lncRNA NARL	miR-217-5p	Sponge effect	[39]
circRNA	circRasGEF1B	miR-21-3p (targeting MITA)	Sponge effect	[50]
	circCBL	miR-125a-1-3p (targeting MITA)	Sponge effect	[51]
	circDtx1	miR-15a-5p (targeting TRIF)	Sponge effect	[52]
	circSamd4a	miR-29a-3p (targeting STING)	Sponge effect	[53]
	circRNF217	miR-130-3p (targeting NOD1)	Sponge effect	[54]
	circBCL2L1	miR-30c-3-3p (targeting TRAF6)	Sponge effect	[55]
	circPIKfyve	miR-21-3p (targeting MAVS)	Sponge effect	[56]
	circVPS13D	VPS13D-170aa (targeting MAVS)	Coding protein	[57]
	circMORC3	MORC3-84aa (targeting TRIF)	Coding protein	[58]
	circYthdc2	Ythdc2-170aa (targeting STING)	Coding protein	[59]
	circNLRP12	NLRP12-119aa (targeting SCRV)	Coding protein	[60]

RNA type

RNA name

Targeting molecule

Function mode

References

miRNA

miR-217

TGF-b-activated kinase 1 (TAK1)

Inhibition of protein expression (IPE)

[38]

miR-217-5p

Nucleotide oligomerization domain 1 (NOD1)