Table 1.
Mechanisms used by several virus types to release from the host cells
| Type of virus | Family | Intracellular transfer | Hijacking mechanism | Releasing mechanisms | Reference |
|---|---|---|---|---|---|
| Enveloped DNA viruses | |||||
| Epstein-Barr Virus | Herpesviridae | ESCRT-dependent and ESCRT-independent pathways | TSG101, CD63, Syntenin‑1, Alix, and lipid rafts | Late endosomes | [195, 196, 238] |
| Kaposi's sarcoma-associated herpesvirus | Herpesviridae | ESCRT-dependent and ESCRT-independent pathways | Viral LMP1 | Late endosomes | [182, 239] |
| Human Herpes virus-6 | Herpesviridae | ESCRT-independent pathway | CD63 and viral gB | Late endosomes | [172] |
| Human Cytomegalovirus | Herpesviridae | ESCRT-independent, Vamp3-dependent pathways | VAMP3, viral pUL71 and UL99 | Late endosomes | [197, 240] |
| Herpes simplex virus 1 | Herpesviridae | ESCRT-dependent pathway | ESCRT-III |
Microvesicular budding, Autophagosome-mediated exit without lysis, Late endosomes |
[241, 242] |
| Non-enveloped DNA viruses | |||||
| Adenoviruses 2 and 5 | Adenoviridae | ESCRT-independent pathway | Exosomal CD46, Lamp2c | Late endosomes, Microvesicular budding | [181, 243, 244] |
| Hepatitis B virus | Hepadnavirideae | ESCRT-dependent pathway | α-taxilin, viral LHBs, TSG101, 2-adaptin, Nedd4 | Late endosomes, Microvesicular budding | [245] |
| Enveloped RNA viruses | |||||
| Japanese encephalitis virus | Flaviviridae | ESCRT-dependent pathway | viral NS3, TSG101 | Late endosomes | [246, 247] |
| Yellow fever virus | Flaviviridae | ESCRT-dependent pathway | Viral NS3 protein, Alix | Late endosomes | [248, 249] |
| Dengue virus | Flaviviridae | ESCRT-dependent pathway | Viral NS3, Alix, viral protein E, Tsp29Fb | Late endosomes | [186, 187, 249, 250] |
| West Nile virus | Flaviviridae | ESCRT-dependent pathway | Viral NS3, Alix and ESCRT | Late endosomes | [249] |
| Zika virus | Flaviviridae | ESCRT-dependent pathway | Viral NS3 protein interacts with Alix | Late endosomes | [249, 251] |
| Tick-borne encephalitis | Flaviviridae | ESCRT-dependent pathway | ND | Late endosomes | [249] |
| Hepatitis C Virus | Flaviviridae | ESCRT-dependent pathway | Viral NS2 and NS5A proteins, Ago2, HSP90, and CD81 | Late endosomes | [252–254] |
| Human immunodeficiency virus-1 | Retroviridiae | ESCRT-dependent pathway | Viral Nef, Alix, viral Gag, and Syntenin | Late endosomes | [161, 191, 255, 256] |
| Equine Infectious Anemia Virus | Retroviridiae | ESCRT-dependent pathway | Viral Gag, Alix |
Late endosomes TGN is the assembly site |
[257] |
|
Feline immunodeficiency virus |
Retroviridiae | ESCRT-dependent pathway | Viral Gag, and Alix | Microvesicular budding | [258, 259] |
| Human immunodeficiency virus-2 | Retroviridiae | ESCRT-dependent pathway | Viral Nef, Alix and viral Gag, and TSG-101 | Late endosomes, Microvesicular budding | [260, 261] |
| Mouse Mammary Tumor Virus | Retroviridiae | ESCRT-dependent pathway | Viral Gag and NEDD4 | ND | [262] |
| Murine Leukemia Virus | Retroviridiae | ESCRT-dependent pathway | Viral Gag and WWP1 | Late endosomes, Microvesicular budding | [260, 263, 264] |
|
Porcine endogenous retrovirus |
Retroviridiae | ESCRT-dependent pathway | Viral Gag, and WWP2 | Late endosomes | [265] |
| Prototypic foamy viruses | Retroviridiae | ESCRT-dependent pathway | Viral Gag, TSG-101, and PPPI | Late endosomes, Microvesicular budding, trans-Golgi network | [266, 267] |
| Mason–Pfizer monkey virus | Retroviridiae | ESCRT-dependent pathway | Viral Gag, TSG-101 and NEDD4 | Microvesicular budding | [268, 269] |
|
Human T cell leukemia virus type 1 |
Retroviridiae | ESCRT-dependent pathway | Viral Gag, TSG-101 and NEDD4 | Late endosomes | [260, 270] |
| Human endogenous retrovirus-K | Retroviridiae | ESCRT-dependent pathway | Viral Gag, TSG-101 | Late endosomes | [271, 272] |
| Rous sarcoma virus | Retroviridiae | ESCRT-dependent pathway | Gag, NEDD4 and Alix | Late endosomes | [273, 274] |
| Respiratory syncytial virus | Pneumoviridae | ESCRT-independent pathway | ND | Late endosomes | [46] |
| Human Pegivirus, GB virus C | Flaviviridae | ESCRT-dependent pathway | hnRNPA2B1 protein | Late endosomes | [275, 276] |
| Ebola virus | Flaviviridae | ESCRT-dependent pathway | Viral VP40, TSG-101, NEDD4, and VP24 | Late endosomes, Microvesicular budding | [277, 278] |
| Marburg virus | Flaviviridae | ESCRT-dependent pathway | TSG-101, Viral VP40 and NEDD4 | Microvesicular budding | [279–281] |
| Lassa fever virus | Arenaviridae | ESCRT-dependent pathway | Z protein and TSG-101 | Microvesicular budding | [282, 283] |
| Rabies Virus | Rhabdoviridae | ESCRT-dependent pathway | Viral M protein, ESCRTI, Alix, ESCRT III and VPS4 | Microvesicular budding | [284, 285] |
| Vesicular stomatitis virus | Rhabdoviridae | ESCRT-dependent pathway | Viral M protein, TSG-101, and NEDD4 | Microvesicular budding | [286] |
| Parainfluenza virus 5 | Paramyxoviridae | ESCRT-dependent pathway | Viral M protein, and VPS4 ATPase | Microvesicular budding | [287] |
| Parainfluenza virus 3 | Paramyxoviridae | ESCRT-dependent pathway | Viral M protein | Microvesicular budding | [288] |
| Newcastle disease | Paramyxoviridae | ESCRT-dependent pathway | Viral M protein and VPS4 ATPase | Microvesicular budding | [289] |
| Mumps virus | Paramyxoviridae | ESCRT-dependent pathway | Viral M protein and VPS4 ATPase | Microvesicular budding | [290] |
| Hepatitis E virus | Hepeviridae | ESCRT-dependent pathway | Viral pORF3, and TSG-101 | Late endosomes | [291, 292] |
| Non-enveloped RNA viruses | |||||
| Hepatitis A virus | Picornaviridae | ESCRT-dependent pathway | Viral VP2, Alix and VPS4B | Late endosomes | [166, 293] |
| Coxsackievirus B | Picornaviridae | ESCRT-dependent pathway | LC3 | Microvesicular budding, Autophagosome-mediated exit without lysis and | [294] |
| Poliovirus | Picornaviridae | ESCRT-dependent and independent pathways | Alix | Microvesicular budding, Autophagosome-mediated exit without lysis and | [165] |
| Enterovirus 71 | Picornaviridae | ESCRT-dependent pathway | ND | Late endosomes | [295–297] |
| Reovirus | Reoviridae | ESCRT-dependent pathway | Rab5, Viral capsid protein P2 | Late endosomes, modified lysosomal organelle act as transporter virions to the plasma membrane | [184, 298] |
| Rotavirus | Reoviridae | Probably ESCRT-dependent pathway | Probably VP4 interacts with lipid Rafts | Microvesicular budding | [150] |
| Norovirus | Caliciviridae | Probably ESCRT-dependent pathway | Viral VP1, Phosphatidylserine lipid | Late endosomes | [174] |