PMC:7272971 / 7922-14415 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/7272971","sourcedb":"PMC","sourceid":"7272971","source_url":"https://www.ncbi.nlm.nih.gov/pmc/7272971","text":"Binding of coronaviruses to host cells, and presence of receptors in ocular surface tissue\nSARS‐CoV‐2 uses angiotensin converting enzyme‐2 (ACE2) as its receptor on human cells, and binding to this facilitates infection. 16 The mammalian serine protease TMPRSS2 or the protease Furin (also known as Paired basic Amino acid Cleaving Enzyme) appear to prime the spike protein for interaction with ACE2.17, 18 SARS‐CoV also binds to the cell‐surface protein ACE2 and this binding is required for the virus to infect cells. 19 The binding of SARS‐CoV‐2 to ACE2 is enhanced compared to that of SARS‐CoV due to several changes in the gene and hence amino acid sequence of the spike protein. 20 SARS‐CoV also binds to CD209 (dendritic cell [DC]‐specific intercellular adhesion molecule [ICAM‐3] grabbing non‐integrin; DC‐SIGN) and CD209L (L‐SIGN).21, 22, 23 MERS‐CoV binds to the cell‐surface molecule CD26 (DPP‐4) and this is facilitated by the cofactor CD66e (CEACAM5). 24\nFor the less pathogenic human coronaviruses, hCoV‐229E uses CD13 to infect cells, 25 OC43 and HKU1 bind to 9‐O‐acetylated sialic acid (Neu5,9Ac2), 26 and NL63 uses ACE2 to bind to host cells. 27 NL63 and SARS‐CoV first anchor to cells via heparan sulfate proteoglycans before interacting with the ACE2 entry receptor,19, 27, 28 but this binding alone is not sufficient for infection. 19 In fact many viruses, not just coronaviruses, utilise heparan sulfate on cell surfaces for initial binding. 29 Feline infectious peritonitis virus uses CD13 (aminopeptidase N) or CD209 to bind to its host cells. 15 Bovine coronavirus binds to 9‐O‐acetylated sialic acid on its host cells. 30\nTable 1 presents a comparison of receptors for SARS‐CoV‐2, SARS‐CoV and MERS‐CoV on ocular surface and lung cells.31, 32, 33, 34, 35, 36, 37, 38, 39\nTable 1 Comparison of SARS‐CoV‐2, SARS‐CoV and MERS‐CoV receptors on ocular surface and lung\nCoronaviruses Receptor Role in human CoV disease Ocular surface (cell types) Lung\nSARS‐CoV\nSARS‐CoV‐2 ACE2 Host cell receptor and essential for infection + (conjunctival epithelium and fibroblasts; corneal epithelium) 31 + (airway epithelia)32, 33\nTMPRSS2 Primes spike protein for binding to ACE2 – + (airway epithelium) 34\nCD209 Cell‐to‐cell viral transfer + (corneal dendritic cells) 35 + (alveola macrophages) 36\nMERS‐CoV CD26 Host cell receptor and essential for infection +/− (conjunctival vascular endothelium) 37 + (lung epithelia and endothelia) 38\nCD66e Co‐receptor +/− (palpebral conjunctiva) 39 + (bronchial and alveolar epithelium) 32\n+: the receptor has been found, −: there are no reports of the receptor on the ocular surface, +/−: found in tissue but only on non‐virally‐mediated inflamed tissue. Expression of the gene for ACE2 has been shown in corneal and conjunctival epithelial cells, and preliminary evidence has been presented for the presence of the protein associated with these cells. 40 The isolated surface protein S240 of SARS‐CoV can bind to conjunctival epithelial and fibroblast cells and corneal epithelial cells, with this binding inhibited by soluble ACE2, indicating that the binding was being mediated through cell‐surface ACE2. 31 However, the authors did note that ‘the amount of ACE2 expressed in conjunctival and corneal cells and tissues (was) less than that in heart, lung tissues and Vero E6 cells’, and was approximately 50 per cent that of Vero E6 cells. 31 Another paper provided evidence of expression of mRNA for ACE2 in corneal epithelium with no evidence provided of mRNA from the conjunctiva. 41 The normalised expression of ACE2 in the corneal epithelial mRNA was only 20‐28 units, compared to \u003e 400 units in the testes, \u003e 200 units in the small intestine terminal ileum and \u003e 100 units in the heart left ventricle. 41 The previously mentioned serine protease TMPRSS2 or Furin, which appear to prime the spike protein, had not been reported in tears or on corneal or conjunctival cells. However, 15 per cent of superficial conjunctival cells have been shown to have “greater‐than‐zero expression” of the gene for TMPRSS2, three per cent of cells in superficial conjunctiva and one per cent of limbal superficial cells have “greater‐than‐zero expression” of genes of both TMPRSS2 and ACE2. The mean level of expression of mRNA for ACE2 or TMPRSS2 in any ocular tissue was approximately 0.6 per cent, whereas the mean level of expression was greater (1.2–1.4 per cent) in most other tissues (such as nasal cavity, lung parenchyma, ileum, colon, heart) examined. 42\nCD209 can be found on human corneal dendritic cells. 35 CD26 is not expressed in normal conjunctival tissue, and has only been found, and then only weakly expressed, in vascular endothelial cells of conjunctival tissue from biopsies of vernal conjunctivitis patients. 37 The only report of CD66e in the conjunctiva was from patients with trachoma who had gene expression in their upper palpebral conjunctiva. 39 There are no reports of CD26 or CD66e in cornea or tears. CD13 has not been reported on conjunctival or corneal epithelial cells, but has been found on human corneal keratocytes. 43 9‐O‐acetylated sialic acid has been found in human tears and on human corneal mucin. 44 Heparan sulfate has been found on human cornea and conjunctiva cells.45, 46\nIn summary, the ocular surface does possess the ubiquitous viral receptor heparan sulfate, which can facilitate initial viral attachment. However, this is not sufficient for coronavirus infection. Furthermore, lactoferrin that is present in high concentrations in tears (2 mg/ml) 47 prevents viral attachment to heparan sulfate. 19 The ocular surface appears to possess the SARS‐CoV‐2, SAR‐CoV and NL63 receptor ACE2, but may not possess the TMPRSS2 or Furin proteins, which is required for the spike protein of SARS‐CoV‐2 to bind to ACE2.\nFurthermore, apart from 9‐O‐acetylated sialic acid, the other co‐receptors for coronaviruses (CD209, CD26, CD13, CD66e), have been found either in fibroblasts and dendritic cells that lie in tissue under the surface corneal and conjunctival epithelial cells, or have not been reported in ocular tissues. It is possible that the 9‐O‐acetylated sialic acid associated with proteins in tears can act as a decoy receptor, binding to the viruses and preventing them binding to cell‐associated 9‐O‐acetylated sialic acid. 44 Therefore, the evidence to date suggests that coronaviruses are unlikely to bind to ocular surface cells to initiate 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