PMC:7103735 / 35504-41775 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/7103735","sourcedb":"PMC","sourceid":"7103735","source_url":"https://www.ncbi.nlm.nih.gov/pmc/7103735","text":"Evolving to be less pathogenic: is there a trend?\nCoVs are believed to have existed for \u003e6000 years. Molecular clock analysis enables us to deduce the time of emergence or cross-species transmission of some HCoVs. Surprisingly, the highly pathogenic SARS-CoV and MERS-CoV are thought to emerge in the last 30 years [56]. Although NL63 and HKU1 of low pathogenicity were isolated in the post-SARS era, their time of emergence was earlier than the other two highly pathogenic viruses. Whereas NL63 emerged \u003e500 years ago [57], the origin of HKU1 dated back to the 1950s. In other words, SARS-CoV and MERS-CoV represent newcomers that are striving to adapt to humans, whereas other community-acquired HCoVs including NL63 and HKU1 are better adapted. The results of molecular dating of HCoVs suggest that attenuation might be favoured during evolution when host adaptation takes place.\nTo shed further light on this trend, a comparative analysis of the innate immunomodulatory activity of viral proteins encoded by different HCoVs would be helpful. In the first place, type I interferon antagonism of these proteins might be compared. The use of interferon to treat HCoV-infected patients was tested as early as the 1960s, when various volunteer experiments were performed to investigate how humans combat HCoV infection. Administration of interferon ameliorated the severity of symptoms in 229E-challenged volunteers [58], suggesting that interferon confers protection against 229E infection. Indeed, 229E potently induces type I interferon and its replication is susceptible to inhibition by type I IFN [59], suggesting that interferon serves as a key component of host antiviral defence. Since interferon effectively suppresses the early phase of viral replication, the suppression of interferon production and signalling by highly pathogenic HCoVs can exacerbate disease progression. Thus, highly pathogenic HCoVs adopt various countermeasures to suppress host interferon production and signalling as reviewed elsewhere [49].\nHost innate immune response is the first-line defence triggered by type I interferon. Type I interferon production is activated through the detection of replicating viral RNA by cytoplasmic RNA sensors RIG-I and MDA5. Oligomerization of adaptor protein MAVS is induced by the activation of RNA sensor, leading to the formation of TRAF3-TANK-TBK1/IKKϵ complex, which phosphorylates transcription factor IRF3 and drives type I IFN transcription [49]. Highly pathogenic HCoVs often encode viral proteins with a higher capability to antagonize RNA-induced type I interferon production through perturbation of RNA sensing. For one example, double-stranded RNA (dsRNA)-binding domain of MERS-CoV ORF4a is responsible for the suppression of Sendai virus- or poly (I:C)-induced type I interferon production [60]. The gain of dsRNA-binding ability is observed in bat CoV HKU5 but not HKU4, suggesting that the functional gain of MERS-CoV ORF4a and HKU5 ORF4a might be acquired at a later stage in evolution [60]. For another example, only M proteins from highly pathogenic CoVs, SARS-CoV and MERS-CoV, were reported to potently suppress type I interferon production [61,62], suggesting that the loss of this activity might have taken place during viral evolution, leading to attenuation.\nOther than suppressing type I interferon induction, both alphacoronaviruses and betacoronaviruses can use nsp1 to degrade host mRNA transcripts, resulting in the suppression of host interferon response, although sequence homology between nsp1 proteins of the two genera remains low. A conserved domain is present in nsp1 of alphacoronaviruses and it is responsible for the shut-down of host gene expression. The deletion of 91–95 amino acids in 229E and NL63 nsp1 partially restores host gene expression as shown by luciferase reporter assay [63]. The interferon-modulating activity of nsp1 shows some variations within the genus of Betacoronavirus. Whereas nsp1 proteins encoded by SARS-CoV and bat CoV Rm1 potently suppress the induction of type I interferon, counterparts in bat CoV 133 and bat CoV HKU9-1 are relatively weak interferon suppressors [64]. Again, some degree of conservation in interferon antagonism of nsp1 is seen among different CoVs.\nApart from preventing interferon production, CoV proteins have also evolved to suppress interferon effector signalling. One of the best-characterized examples is the oligoadenylate synthetase (OAS)-RNase L pathway. Upon activation by interferon in response to the sensing of dsRNA, transcription and expression of OAS are induced to catalyse 2′,5′-oligoadenylate (2′-5′A) synthesis [65]. 2′-5′A serves as a second messenger, which activates RNase L and limits viral replication through the cleavage of cellular and viral single-stranded RNA [65]. The cleaved RNA fragments, in turn, prime RNA sensors to amplify interferon production in infected cells. To counter RNase L activity, some CoVs have evolved phosphodiesterases which cleave 2′-5′A [66]. The first CoV phosphodiesterase identified is NS2a in mouse hepatitis virus. From protein sequence alignment, multiple CoVs including NS2a from OC43 and a bat CoV, as well as ORF4b encoded by MERS-CoV, are very similar to NS2a of mouse hepatitis virus. All these viral proteins preserve a phosphodiesterase activity [67]. This suggests that some CoVs have developed an important enzymatic activity through convergence and divergence. Hence, although most HCoVs retain one or another strategy to counter host antiviral defence, the highly pathogenic CoVs are particularly powerful in the suppression of host immunity. These capabilities might be weakened or lost when they adapt to humans.\nAlthough the exact mechanisms by which SARS-CoV-2 might counteract host antiviral defence remain to be elucidated, this emerging virus is thought to be less pathogenic than SARS-CoV. Our prediction is that the IFN antagonism of SARS-CoV-2 and its ability to suppress other pathways of innate antiviral signalling might fall between those of SARS-CoV and the community-acquired HCoVs. Plausibly, the weakened IFN antagonism of SARS-CoV-2 compared to that of SARS-CoV might lead to more robust host antiviral defence, attenuated viral replication, and lower 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