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Anti-Classical Swine Fever Virus Strategies

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Anti-Classical Swine Fever Virus Strategies microorganisms Review Anti-Classical Swine Fever Virus Strategies Jindai Fan 1,2,3,†, Yingxin Liao 1,2,3,†, Mengru Zhang 1,2,3, Chenchen Liu 1,2,3, Zhaoyao Li 1,2,3, Yuwan Li 1,2,3, Xiaowen Li 1,2,3, Keke Wu 1,2,3, Lin Yi 1,2,3, Hongxing Ding 1,2,3, Mingqiu Zhao 1,2,3, Shuangqi Fan 1,2,3,* and Jinding Chen 1,2,3,* 1 College of Veterinary Medicine, South China Agricultural University, Guangzhou 510642, China; [email protected] (J.F.); [email protected] (Y.L.); [email protected] (M.Z.); [email protected] (C.L.); [email protected] (Z.L.); [email protected] (Y.L.); [email protected] (X.L.); [email protected] (K.W.); [email protected] (L.Y.); [email protected] (H.D.); [email protected] (M.Z.) 2 Guangdong Laboratory for Lingnan Modern Agriculture, College of Veterinary Medicine, South China Agricultural University, Guangzhou 510642, China 3 Key Laboratory of Zoonosis Prevention and Control of Guangdong Province, Guangzhou 510642, China * Correspondence: [email protected] (S.F.); [email protected] (J.C.); Tel.: +86-20-8528-8017 (J.C.) † These authors contributed equally to this work. Abstract: Classical swine fever (CSF), caused by CSF virus (CSFV), is a highly contagious swine disease with high morbidity and mortality, which has caused significant economic losses to the pig industry worldwide. Biosecurity measures and vaccination are the main methods for prevention and control of CSF since no specific drug is available for the effective treatment of CSF. Although a series of biosecurity and vaccination strategies have been developed to curb the outbreak events, it is still difficult to eliminate CSF in CSF-endemic and re-emerging areas. Thus, in addition to implementing enhanced biosecurity measures and exploring more effective CSF vaccines, other strategies are also Citation: Fan, J.; Liao, Y.; Zhang, M.; needed for effectively controlling CSF. Currently, more and more research about anti-CSFV strategies Liu, C.; Li, Z.; Li, Y.; Li, X.; Wu, K.; was carried out by scientists, because of the great prospects and value of anti-CSFV strategies in Yi, L.; Ding, H.; et al. Anti-Classical the prevention and control of CSF. Additionally, studies on anti-CSFV strategies could be used as a Swine Fever Virus Strategies. Microorganisms 2021, 9, 761. https:// reference for other viruses in the Flaviviridae family, such as hepatitis C virus, dengue virus, and doi.org/10.3390/microorganisms Zika virus. In this review, we aim to summarize the research on anti-CSFV strategies. In detail, host 9040761 proteins affecting CSFV replication, drug candidates with anti-CSFV effects, and RNA interference (RNAi) targeting CSFV viral genes were mentioned and the possible mechanisms related to anti-CSFV Academic Editor: Keivan Zandi effects were also summarized. Received: 2 March 2021 Keywords: classical swine fever virus; host protein; antiviral target; antiviral drug; RNA interference Accepted: 2 April 2021 Published: 6 April 2021 Publisher’s Note: MDPI stays neutral 1. Introduction with regard to jurisdictional claims in Classical swine fever (CSF) is a highly contagious swine disease characterized by published maps and institutional affil- high fever, multiple hemorrhages, and gastrointestinal symptoms with high morbidity and iations. mortality, which has caused significant economic losses to the pig industry worldwide [1]. CSF in wild boars is also endemic in some countries, which poses a serious threat to domestic pigs [2]. The causative agent of CSF is the CSF virus (CSFV), an important member of the Pestivirus genus within the Flaviviridae family. CSFV is an enveloped, Copyright: © 2021 by the authors. single-stranded, positive-sense RNA virus containing a 12.3 kb RNA genome, which Licensee MDPI, Basel, Switzerland. consists of a 50-untranslated region (50-UTR), a single large open reading frame (ORF), and This article is an open access article a 30-UTR. A polypeptide precursor could be encoded by ORF of CSFV and then cleaved distributed under the terms and into four structural proteins (C, Erns, E1, and E2) and eight non-structural proteins (Npro, conditions of the Creative Commons p7, NS2, NS3, NS4A, NS4B, NS5A, and NS5B) through the processing of the polypeptide Attribution (CC BY) license (https:// creativecommons.org/licenses/by/ precursor by viral and cellular proteases (Figure1)[1,3,4]. 4.0/). Microorganisms 2021, 9, 761. https://doi.org/10.3390/microorganisms9040761 https://www.mdpi.com/journal/microorganisms Microorganisms 2021, 9, x FOR PEER REVIEW 2 of 17 Microorganisms 2021, 9, 761 2 of 17 FigureFigure 1.1. SchematicSchematic description description of of virion virion structure structure and and genome genome organization organization of classical of classical swine swine fever fevervirus virus(CSFV) (CSFV) [4]. This [4]. figure This figure comes comes from the from literature the literature reported reported by Beer by et Beer al. [4] et with al. [4 some] with modifi- some modifications.cations. CSFVCSFV is an isenveloped, an enveloped, single-strande single-stranded,d, positive-sense positive-sense RNA RNA virus. virus. Glycoproteins Glycoproteins (Erns (E,rns E1,, E1,and and E2) E2)are arelocated located on the on theexternal external part partof viral of viral particles particles and are and important are important for viral for infection. viral infection. CSFV RNA genome consists of a single large open reading frame (ORF) flanked by a 5′-untranslated re- CSFV RNA genome consists of a single large open reading frame (ORF) flanked by a 50-untranslated gion (5′-UTR) and a 3′-UTR. A polypeptide precursor could be encoded by ORF of CSFV and then region (50-UTR) and a 30-UTR. A polypeptide precursor could be encoded by ORF of CSFV and then cleaved into four structural proteins (C, Erns, E1, and E2) and eight non-structural proteins (Npro, p7, rns pro cleavedNS2, NS3, into NS4A, four structural NS4B, NS5A, proteins and (C,NS5B). E , E1, and E2) and eight non-structural proteins (N , p7, NS2, NS3, NS4A, NS4B, NS5A, and NS5B). BasedBased onon partialpartial sequencessequences ofof CSFVCSFV 550-UTR,′-UTR, E2,E2, andand NS5B,NS5B, phylogenicphylogenic typingtyping ofof CSFVCSFV isolates isolates worldwide worldwide was was carried carried out. out. In In general, general, CSFV CSFV strains strains could could be be classified classified into into threethree genotypes genotypes (1, (1, 2, 2, and and 3) 3) and and eleven eleven subgenotypes subgenotypes (1.1–1.4, (1.1-1.4, 2.1–2.3, 2.1-2.3, and and 3.1–3.4) 3.1-3.4) [[5,6];5,6]; subgenotypesubgenotype 2.1 2.1 isolates isolates were were further further divided divided into into three three sub-subgenotypes sub-subgenotypes (2.1a–2.1c) (2.1a–2.1c) [7– [7–9]. Because9]. Because of the of highthe high genetic genetic diversity diversity of subgenotype of subgenotype 2.1 strains, 2.1 strains, Gong Gong et al. [10et ]al. suggested [10] sug- dividinggested dividing the subgenotype the subgenotype 2.1 strains 2.1 intostrains ten in sub-subgenotypesto ten sub-subgenotypes (2.1a–2.1j). (2.1a–2.1j). As reported, As re- CSFVported, strains CSFV of strains genotype of genotype 1, 2, and 31, are 2, and all epidemic 3 are all inepidemic the world, in the while world, strains while of genotype strains of 2genotype have gradually 2 have becomegradually dominant become globallydominant [11 globally–14]. [11–14]. CSFVCSFV strains’ strains’ virulence, virulence, ranging ranging from from high, high, moderate, moderate, to to low low virulence, virulence, is is the the crucial crucial determinantdeterminant of of CSFV CSFV pathogenicity pathogenicity in in pigs. pigs. CSFVCSFV infectioninfection cancan leadlead toto anan acute,acute, subacute,subacute, chronic,chronic, or or asymptomatic asymptomatic disease disease in in pigs pigs depending depending on on CSFV CSFV strains’ strains’ virulence virulence and and other other factors,factors, such such as as the the host’s host’s age age and and immune immune status status [15 [15].]. Although Although CSFV CSFV strains strains with with different differ- virulenceent virulence exist exist globally, globally, more andmore more and reportsmore reports show that show the that currently the currently circulating circulating strains arestrains mainly are moderatelymainly moderately virulent virulent [16–19]. [16–19]. Currently,Currently, the the World World Organization Organization for for Animal Anim Healthal Health (OIE) (OIE) lists lists 38 CSF-free 38 CSF-free members, mem- whichbers, which mainly mainly locate locate in North in North America, America, part part of Southof South America, America, Oceania, Oceania, and and a a large large partpart ofof the European Union. Union. CSF CSF is isstill still endemic endemic in Asia, in Asia, South South and andCentral Central America, America, East- Easternern Europe, Europe, and and parts parts of Africa. of Africa. Additionally, Additionally, specific specific zones zones in Brazil, in Brazil, Colombia, Colombia, and and Ec- Ecuadoruador are are declared declared CSF-free CSF-free by byOIE, OIE, while while the theother other zones zones in these in these countries countries remain remain CSF- CSF-endemicendemic [20]. [ 20Additionally,]. Additionally, there there is a isrisk a risk of CSF of CSF re-emergence re-emergence in inCSF-free CSF-free areas areas due due to tothe the existence existence of ofCSF-endemic CSF-endemic regions regions in the in the world, world, as well as well as the as thereservoir reservoir of CSFV of CSFV in wild in wildboars. boars. For Forinstance, instance, in 2018, in 2018, CSF CSF re-emerged re-emerged in inJapan Japan after after 26 26 years years of CSF-free status,status, affectingaffecting bothboth domesticdomestic pigspigs andand wildwild boarsboars [ 12[12].].
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