Table 1.
Interaction of heparan sulfate and SARS-CoV-2
| References | Role of Heparan Sulfate | The Types of Virus Stains/Protein | Note |
|---|---|---|---|
| Clausen et al. (14) | Attachment factor | 1) SARS-CoV-2 live virus (USA-WA1/2020)2) pp-SARS2 | Binding to HS changes the conformation of the spike protein |
| Kim et al. (83) | Interaction with recombinant S | Recombinant SARS-CoV-2 spike protein | Recombinant spike protein interacts with the GAG-binding motif at the S1/S2 site and [453-459 (YRLFRKS)] |
| Mycroft-West et al. (12) | Heparin inhibits the SARS-CoV-2 entry | Italy/UniSR1/2020 isolate (GISAID accession ID: EPI_ISL_413489) | Binding to the RBD is more likely dependent on the structures of 2-O- or 6-O-sulfate groups |
| Schuurs et al. (84) | Interaction with spike protein | Bioinformatics and molecular dynamics simulations | The heparan sulfate binds the furin cleavage site and S247R; heparin binds the furin cleavage site and surrounding glycosylation structures, but not S247R |
| Chu et al. (10) | Attachment factor | 1) SARS-CoV-2 HKU-001a (GenBank accession number MT230904)2) SARS-CoV-2 HKU-001a S1/S2mut (GenBank accession number MT621560)3) pp-SARS2 | Both human lung epithelial cells and ex vivo human lung tissues were used in assessing the role of heparan sulfate |
| Tandon et al. (85) | Sulfated polysaccharides exhibit potent anti-SARS-CoV-2 activity | pp-SARS2 | |
| Yan et al. (86) | Binding to the spike protein | pp-SARS2 | SARS-CoV-2 spike protein binds stronger to human lung heparan sulfate than bat lung heparan sulfate |
| Partridge et al. (87) | Unfractionated heparin inhibits spike protein interaction | Recombinant SARS-CoV-2 spike protein | Chondroitin sulfate did not inhibit the binding of the spike protein |
| Yue et al. (88) | Facilitates spike protein-mediated SARS-CoV-2 cell entry | pp-SARS2 | Overexpression of Ext1 in 293 T-ACE2 cells increased pp-SARS2 infection |
| Liu et al. (89) | Attachment factor | Recombinant protein | HS Functions as initial attachment factor, which facilitates the binding of spike protein to ACE2 |
| Hao et al. (11) | Binding to the spike protein | Recombinant protein | S proteins are able to bind to HS in a sulfation-dependent manner specifically |
| Gupta et al. (90) | Heparin inhibits SARS-CoV-2 infection | SARS-COV2/MT020880.1 | |
| Shiliaev et al. (91) | Heparin can effectively inhibit Cell adapted SARS-CoV-2 but not natural isolates | 1) SARS-CoV-2 strain 2019nCoV/USA_WA1/20202) CoV-2/UAB | Serial passages of natural SARS-CoV-2 isolates in Vero-E6 increases the affinity to HS |
| Bermejo-Jambrina et al. (92) | Attachment receptor | 1) SARS-Related Coronavirus 2, Isolate Italy-INMI1, NR-52284; 2) PP-SARS2 | Heparan sulfate proteoglycans Syndecan 1 and 4 are important for SARS-CoV-2 binding |
| Chittum et al. (93) | Interacts with SARS-CoV-2 spike | HS microarray/SARS-CoV-2 protein | Equivalent importance of IdoA2S−GlcNS6S and GlcNS3S on binging to RBD |
| Zheng et al. (94) | Heparin binding to SARS-CoV-2 spike protein | mFc-tagged S1 protein | Spike protein directly promotes blood coagulation and thrombosis in the zebrafish model |
HS, heparin sulfate; pp-SARS2: pseudotyped SARS-CoV-2.