key: cord-293082-fw7deem8
authors: Zhang, Guangzhi; Li, Bin; Yoo, Dongwan; Qin, Tong; Zhang, Xiaodon; Jia, Yaxiong; Cui, Shangjin
title: Animal coronaviruses and SARS‐CoV‐2
date: 2020-08-16
journal: Transbound Emerg Dis
DOI: 10.1111/tbed.13791
sha: 
doc_id: 293082
cord_uid: fw7deem8

COVID‐19 is a highly contagious disease caused by severe acute respiratory syndrome coronavirus 2 (SARS‐CoV‐2). It has rapidly spread to 216 countries and territories since first outbreak in December of 2019, posing a substantial economic losses and extraordinary threats to the public health worldwide. Although bats have been suggested as the natural host of SARS‐CoV‐2, transmission chains of this virus, role of animals during cross‐species transmission, and future concerns remain unclear. Diverse animal coronaviruses have extensively been studied since the discovery of avian coronavirus in 1930s. The current article comprehensively reviews and discusses the current understanding about animal coronaviruses and SARS‐CoV‐2 for their emergence, transmission, zoonotic potential, alteration of tissue/host tropism, evolution, status of vaccines, and surveillance. This study aims at providing guidance for control of COVID‐19 and preventative strategies for possible future outbreaks of zoonotic coronavirus via cross‐species transmission.

The coronavirus disease 2019 broke out in Wuhan, China in December 2019, and 46 spread rapidly across the world. On March 11th of 2020, World Health Organization (WHO) 47 announced COVID-19 a pandemic. As of April 7, just four months since its first outbreak, more 48 than 3.4 million confirmed cases and 238,000 deaths have been recorded in 215 countries, areas, 49 and territories, and moreover it seems that severe acute respiratory syndrome coronavirus 2 50 (SARS-CoV-2) that causes COVID-19 will probably continue to circulate around the globe 51 (https://www.who.int/emergencies/diseases/novel-coronavirus-2019/situation-reports/). The health 52 authorities and governments of affected countries have paid attention to current pandemic and 53 have taken immediate measures to block COVID-19 transmission, including utilization of personal 54 protective equipment, quarantine, epidemiological investigation, isolation, clinical data analysis 55 and sharing, public health education, maintaining social distance, the creation of diagnostics, 56 therapeutics, and vaccines, etc (Xiao and Torok 2020) . The new information is rapidly 57 accumulating and uncovers the nature of SARS-CoV-2, but many questions remain to be 58 answered. (Table 1) . These six CoVs usually cause infections in pigs, but PDCoV seems to have the ability 141 to infect other species such as badgers, calves, and cats (Li et al. 2018) . Moreover, it was reported Our phylogenetic analysis with S protein shows that PEDV and TGEV share only 42.8%-43.5% of 154 genome similarity with SARS-CoV-2, and PHEV and PDCoV share 49.2%-49.3% and 155 40.3%-40.4% genome similarity with SARS-CoV-2, respectively (Table 2) . Obviously, porcine

CoVs are unlikely the origin of SARS-CoV-2. However, the RBD of SARS-CoV-2 is likely to 157 recognize porcine ACE2 based on the high similarity of critical virus-binding residues between 158 porcine ACE2 and human ACE2 (Wan, Shang, Graham, et al. 2020 

This article is protected by copyright. All rights reserved

The investigation needs to be explored. IBV normally binds to cell receptors via sialic acid for its 185 attachment and entry (Shahwan et al. 2013; Toro, van Santen, and Jackwood 2012) (Table 1) .

Live-attenuated vaccines for IBV are usually adopted by the farms, which are developed by serial 187 passages of virulent strains in embryonated chicken eggs (Laconi et al. 2020; Masoudi, Pishraft 188 Sabet, and Shahsavadi 2020; Baron, Iqbal, and Nair 2018 

Kong. An OIE report shows that the antibody in a pet dog living with a receptor for both FECV and FIPV (Hohdatsu et al. 1998 ) ( Table 1 ). As shown in Table 2 showed that the receptor-binding motif of SARS-CoV-2 RBD can engage with ACE2 of humans 260 and cats at similar efficiencies (Wan, Shang, Graham, et al. 2020 for neonates through colostrum (Nemoto et al. 2017; Kanno et al. 2013) . 

As for the receptors utilized by SARS-CoV-2, biophysical and structural data showed that the S 374 protein engages with ACE2 with at least 10 times higher affinity compared to that of SARS-CoV 375 (Wrapp et al. 2020 ). In addition to ACE2, SARS-CoV-2 may also invade the cell via another 376 receptor: CD147 Ibrahim et al. 2020 Accepted Article Ibrahim et al. 2020; Hao et al. 2020) . It was reported that Furin is involved in the cleavage of 381 SARS-CoV-2 S protein and further facilitate viral entry . In principle, tissues 382 with consistently high expression of ACE2 or CD147 are likely susceptible to SARS-CoV-2, 383 including small intestine, kidney (renal tubular cells), testis (Leydig cells and cells in seminiferous 384 ducts), gall bladder, heart, stomach, and liver (Fagerberg et al. 2014 ). According to a recent report Shen et al. 2020 

Phylogenetic analysis by researchers from Institut Pasteur in Paris showed that SARS-CoV-2 409 might already circulated in France prior to first local case (Gámbaro et al. 2020 ). All of these data 410 undoubtedly showed an intricate situation of this virus. Thus, more surveillance needs to be taken 411 to explore the origin and diversity of this virus worldwide.

Importantly, the careful analyses demonstrate that some of the animal CoVs have evolved to All data generated or analyzed in this study are included in this manuscript and in the 526 supplementary material.

This article is protected by copyright. All rights reserved 

This article is protected by copyright. All rights reserved 

This article is protected by copyright. All rights reserved 

This article is protected by copyright. All rights reserved strength of infectivity. 

survey of SARS-CoV-2 for experimental, domestic, companion and wild animals excludes intermediate hosts of 35 different species of animals

Human Kidney is a Target for Novel Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2) Infection

Isolation and sequence analysis of canine respiratory coronavirus

Detection of a group 2 coronavirus in dogs with canine infectious respiratory disease

Analysis of the human tissue-specific expression by genome-wide integration of transcriptomics and antibody-based proteomics

Diagnosis of Feline Infectious Peritonitis: A Review of the Current Literature

Phylogenetic network analysis of SARS-CoV-2 genomes

Evolution of infectious bronchitis virus in the field after homologous vaccination introduction', Vet This article is protected by copyright. All rights reserved J. Saif

Feline coronavirus type II strains 79-1683 and 79-1146 originate from a double recombination between feline coronavirus type I and canine coronavirus

Differences in virus receptor for type I and type II feline infectious peritonitis virus

Engineering a Live Attenuated Porcine Epidemic Diarrhea Virus Vaccine Candidate via Inactivation of the Viral 2'-O-Methyltransferase and the Endocytosis Signal of the Spike Protein

Coronavirus Spike Protein and Tropism Changes'

Human coronaviruses OC43 and HKU1 bind to 9-O-acetylated sialic acids via a conserved receptor-binding site in spike protein domain A

COVID-19 spike-host cell receptor GRP78 binding site prediction

Experimental bovine coronavirus in turkey poults and young chickens

Tracking Changes in SARS-CoV-2 Spike: Evidence that D614G Increases Infectivity of the COVID-19 Virus

Triple-reassortant influenza A virus with H3 of human seasonal origin, NA of swine origin, and internal A(H1N1) pandemic 2009 genes is established in Danish pigs

Attenuated live infectious bronchitis virus QX vaccine disseminates slowly to target organs distant from the site of inoculation

Identifying SARS-CoV-2-related coronaviruses in Malayan pangolins

Structure, Function, and Evolution of Coronavirus Spike Proteins

Broad receptor engagement of an emerging global coronavirus may potentiate its diverse cross-species transmissibility

Cats under the shadow of the SARS-CoV-2 pandemic

Emergence of SARS-CoV-2 through recombination and strong purifying selection

Emerging lethal infectious bronchitis coronavirus variants with multiorgan tropism', Transbound Emerg This article is protected by copyright. All rights reserved ferrets

Immunogenicity and Efficacy of Live Infectious Bronchitis 793/B.08IR Vaccine in SPF Chickens

Origins of the 2009 H1N1 influenza pandemic in swine in Mexico

Contribution of the porcine aminopeptidase N (CD13) receptor density to porcine epidemic diarrhea virus infection

Antibody response to equine coronavirus in horses inoculated with a bovine coronavirus vaccine

Emergence and pandemic potential of swine-origin H1N1 influenza virus

Swine enteric coronavirus disease: A review of 4 years with porcine epidemic diarrhoea virus and porcine deltacoronavirus in the United States and Canada

Mutation of the S and 3c genes in genomes of feline coronaviruses

Prevalence of feline coronavirus (FCoV) and feline leukemia virus (FeLV) in Turkish cats

This article is protected by copyright. All rights reserved antibody-binding epitopes in the spike 2 domain and the nucleocapsid protein of feline infectious peritonitis virus

Sialic acid binding properties of soluble coronavirus spike (S1) proteins: differences between infectious bronchitis virus and transmissible gastroenteritis virus

Cell entry mechanisms of SARS-CoV-2

Pathways of cross-species transmission of synthetically reconstructed zoonotic severe acute respiratory syndrome coronavirus

Genomic diversity of SARS-CoV-2

SARS-CoV-2 and SARS-CoV Spike-RBD Structure and Receptor Binding Comparison and Potential Implications on Neutralizing Antibody and Vaccine Development

Canine Respiratory Coronavirus, Bovine Coronavirus, and Human Coronavirus OC43: Receptors and Attachment Factors', Viruses

Prevalence of feline coronavirus antibodies in Japanese domestic cats during the past decade

On the origin and continuing evolution Accepted Article This article is protected by copyright. All rights reserved of SARS-CoV-2

Genetic diversity and selection regulates evolution of infectious bronchitis virus

Structural basis for human coronavirus attachment to sialic acid receptors

Mutational analysis of aminopeptidase N, a receptor for several group 1 coronaviruses, identifies key determinants of viral host range

Early death after feline infectious peritonitis virus challenge due to recombinant vaccinia virus immunization

Feline infectious peritonitis viruses arise by mutation from endemic feline enteric coronaviruses

Pathogenic characteristics of persistent feline enteric coronavirus infection in cats

Receptor Recognition by the Novel Coronavirus from Wuhan: an Analysis Based on Decade-Long Structural Studies of SARS Coronavirus

Molecular Mechanism for Antibody-Dependent Enhancement of Coronavirus Entry

Development of an Inactivated Vaccine Candidate

SARS-CoV-2 invades host cells via a novel route: CD147-spike protein

Importance of Neutralizing Monoclonal Antibodies Targeting Multiple Antigenic Sites on the Middle East Respiratory Syndrome Coronavirus Spike Glycoprotein To Avoid Neutralization Escape

Emerging and re-emerging coronaviruses in pigs

An outbreak of feline infectious peritonitis in a Taiwanese shelter: epidemiologic and molecular evidence for horizontal transmission of a novel type II feline coronavirus

Evidence of recombination in coronaviruses implicating pangolin origins of nCoV-2019

Discovery of seven novel Mammalian and avian coronaviruses in the genus deltacoronavirus supports bat coronaviruses as the gene source of alphacoronavirus and betacoronavirus and avian coronaviruses as the gene source of gammacoronavirus and deltacoronavirus

Cryo-EM structure of the 2019-nCoV spike in the prefusion conformation

Wenjie Tan, and Genhong Cheng. 2020. 'Mutations

Evidence for gastrointestinal infection of SARS-CoV-2

Isolation and Characterization of 2019-nCoV-like Coronavirus from Malayan Pangolins

Taking the right measures to control COVID-19

Systematic Comparison of Two Animal-to-Human Transmitted Human Coronaviruses: SARS-CoV-2 and SARS-CoV

Clinical course and outcomes of critically ill patients with SARS-CoV-2 pneumonia in Wuhan, China: a single-centered, retrospective

Broad Cross-Species Infection of Cultured Cells by Bat HKU2-Related Swine Acute Diarrhea Syndrome Coronavirus and Identification of Its Replication in Murine Dendritic Cells In Vivo Highlight Its Potential for Diverse Interspecies Transmission

2019 novel coronavirus is undergoing active recombination

COVID-19 and veterinarians for one health, zoonotic-and reverse-zoonotic transmissions

and Team Singapore Novel Coronavirus Outbreak Research. 2020. 'Epidemiologic Features and Clinical Course of Patients Infected With SARS-CoV-2 in

SARS-CoV-2 Spike protein variant D614G increases infectivity and retains sensitivity to antibodies that target the receptor binding domain

The D614G mutation in the SARS-CoV-2 spike protein reduces S1 shedding and increases infectivity

SARS-CoV-2 neutralizing serum antibodies in cats: a serological investigation

Probable Pangolin Origin of SARS-CoV-2 Associated with the COVID-19 Outbreak

Biological and genetic characterization of a hemagglutinating coronavirus isolated from a diarrhoeic child

Tissue tropisms opt for transmissible reassortants during avian and swine influenza A virus co-infection in swine

A Genomic Perspective on the Origin and Emergence of SARS-CoV-2

Evolution of infectious bronchitis virus in China over the past two decades

Fatal swine acute diarrhoea syndrome caused by an HKU2-related coronavirus of bat origin

Ophthalmologic evidence against the interpersonal transmission of 2019 novel coronavirus through conjunctiva', medRxiv: Accepted Article This article is protected by copyright

Immunogenicity and safety of a recombinant adenovirus type-5-vectored COVID-19 vaccine in healthy adults aged 18 years or older: a randomised

Contribution of porcine aminopeptidase N to porcine deltacoronavirus infection

Virulence factors in porcine coronaviruses and vaccine design

This article is protected by copyright. All rights reserved

This article is protected by copyright. All rights reserved