Viral budding via the host ESCRT complexes


Many enveloped viruses recruit components of the cellular ESCRT (endosomal sorting complex required for transport) system to mediate host-assisted viral budding .

ESCRT complexes are normally used by the cell for biological functions involving membrane remodeling, such as intraluminal vesicle formation, autophagy or terminal stages of cytokinesis . The ESCRT family consists of ESCRT-0, ESCRT-I, ESCRT-II which are primarily involved in cargo sorting and membrane deformation, and ESCRT-III which cleaves the bud neck from its cytosolic face . In the last step, vps4 disassembles the complex. The budding reaction catalyzed by the ESCRT machinery has reversed topology when compared with most other budding processes in the cell, such as endocytosis and formation of transport vesicles.



Specific motifs in viral proteins called late assembly domains (L-domains) are responsible for recruiting the host ESCRT machinery directly or via ESCRT-associated proteins like ALIX.

- PTAP motif (and PSAP) recruits the ESCRT machinery via Tsg101.

- LYPx nL motif bind the adaptor protein ALIX, which in turn binds and recruits the ESCRT machinery.

- PPxY motif enlists members of the NEDD4 family of ubiquitin ligases through interactions with WW domains, although the precise link between the NEDD4-like proteins and the ESCRT machinery is still unknown.

- A fourth motif encoded by FPIV has been identified in the matrix protein of the paramyxovirus simian virus 5.
Some enveloped viruses however seem to use other mechanisms to bud, as it has been shown for Influenza virus .

Viruses budding through the ESCRT machinery:

Family Virus Viral protein L-domain motif Host ESCRT Budding location ref
Retroviridae HIV-1 P6-gag PTAP TSG101 Cell membrane
LYPx nL ALIX (low affinity) Cell membrane
SIV P6-gag PTAP TSG101 Cell membrane
LYPx nL ALIX Cell membrane
HTLV-1 P19-gag PPxY NEDD4 Cell membrane
PTAP TSG101 (low affinity) Cell membrane
Human spumaretrovirus Gag protein PTAP TSG101 Cell membrane
Moloney murine leukemia virus P12-gag PPxY ITCH Cell membrane
PTAP TSG101 Cell membrane
LYPx nL ALIX Cell membrane
Bovine leukemia virus Gag protein PPxY nL NEDD4 Cell membrane
Rous sarcoma virus Gag protein PPxY NEDD4 Cell membrane
Mason-Pfizer monkey virus Gag protein PSAP TSG101 Cell membrane
PPxY NEDD4 Cell membrane
EIAV P9-gag LYPx nL ALIX Cell membrane
Hepadnaviridae HBV ?Capsid protein ? ALIX Endoplasmic reticulum
Filoviridae Ebolavirus Matrix protein PTAP TSG101 Cell membrane
PPxY NEDD4 Cell membrane
Marburgvirus Matrix protein PPxY NEDD4 Cell membrane
Nucleoprotein PSAP TSG101 Cell membrane
Rhabdoviridae Vesicular stomatitis virus M protein PPxY NEDD4 Cell membrane
?PSAP ? Cell membrane
Rabies virus M protein PPxY ? Cell membrane
Paramyxoviridae Sendai virus M protein YLDL ALIX Cell membrane
Nipah virus M protein YMYL ? Cell membrane
Simian virus 5 M protein FPIV ? Cell membrane
Arenaviridae Lymphocytic choriomeningitis virus Z protein PTAP TSG101 Cell membrane
Lassa virus Z protein PTAP TSG101 Cell membrane
PPxY ? Cell membrane
Bunyaviridae Gn glycoproteins YxxL? ? Golgi
Flaviviridae Hepatitis C virus ?Core protein ? ? Endoplasmic reticulum
Herpesviridae Herpes simplex virus 1 ? ? Vps4 Trans-Golgi
Epstein-Barr virus ? ? ? Trans-Golgi
HCMV ? ? Vps4 Trans-Golgi
Poxviridae Vaccinia virus F13 protein LYPx(n)L ? Cell membrane

Matching UniProtKB/Swiss-Prot entries

139 entries grouped by protein (browse by keywords)

Only complete proteome entries are shown (view all 207 entries)

4 entries

Envelope protein F13 (37 kDa protein) (Palmitoylated EV membrane protein) (p37K)

Select_allDeselect_all
F13_VACCC
Vaccinia virus (strain Copenhagen) (VACV) reference strain
F13_VACCW
Vaccinia virus (strain Western Reserve) (VACV) (Vaccinia virus (strain WR)) reference strain
F13_VAR67
Variola virus (isolate Human/India/Ind3/1967) (VARV) (Smallpox virus) reference strain
F13_VACCA
Vaccinia virus (strain Ankara) (VACV)

63 entries

Gag polyprotein (Pr55Gag)

Select_allDeselect_all
GAG_FOAMV
Human spumaretrovirus (SFVcpz(hu)) (Human foamy virus) reference strain

Gag polyprotein (Pr71Gag)

GAG_HV190
Human immunodeficiency virus type 1 group M subtype H (isolate 90CF056) (HIV-1) reference strain
GAG_HV1B1
Human immunodeficiency virus type 1 group M subtype B (isolate BH10) (HIV-1) reference strain
GAG_HV1EL
Human immunodeficiency virus type 1 group M subtype D (isolate ELI) (HIV-1) reference strain
GAG_HV1H2
Human immunodeficiency virus type 1 group M subtype B (isolate HXB2) (HIV-1) reference strain
GAG_HV1MA
Human immunodeficiency virus type 1 group M subtype A (isolate MAL) (HIV-1) reference strain
GAG_HV1MN
Human immunodeficiency virus type 1 group M subtype B (isolate MN) (HIV-1) reference strain
GAG_HV1MV
Human immunodeficiency virus type 1 group O (isolate MVP5180) (HIV-1) reference strain
GAG_HV1RH
Human immunodeficiency virus type 1 group M subtype B (isolate RF/HAT3) (HIV-1) reference strain
GAG_HV1VI
Human immunodeficiency virus type 1 group M subtype F1 (isolate VI850) (HIV-1) reference strain
GAG_HV1YF
Human immunodeficiency virus type 1 group N (isolate YBF30) (HIV-1) reference strain
GAG_HV2BE
Human immunodeficiency virus type 2 subtype A (isolate BEN) (HIV-2) reference strain
GAG_MLVMS
Moloney murine leukemia virus (isolate Shinnick) (MoMLV) reference strain

Gag polyprotein (Pr65gag) (Core polyprotein)

GAG_MMTVC
Mouse mammary tumor virus (strain C3H) (MMTV) reference strain

Gag polyprotein

GAG_SFV1
Simian foamy virus type 1 (SFVmac) (SFV-1) reference strain

Gag polyprotein (Pr71Gag)

GAG_SFVCP
Simian foamy virus (isolate chimpanzee) (SFVcpz) reference strain

Gag polyprotein (Pr71Gag)

GAG_SIVEK
Simian immunodeficiency virus (isolate EK505) (SIV-cpz) (Chimpanzee immunodeficiency virus) reference strain
GAG_SIVMB
Simian immunodeficiency virus (isolate MB66) (SIV-cpz) (Chimpanzee immunodeficiency virus) reference strain
GAG_SIVTN
Simian immunodeficiency virus (isolate TAN1) (SIV-cpz) (Chimpanzee immunodeficiency virus) reference strain
GAG_XMRV6
Xenotropic MuLV-related virus (isolate VP62) (XMRV) reference strain

Gag polyprotein (Pr65gag) (Core polyprotein)

GAG_AVISY
Avian sarcoma virus (strain Y73)

Gag-yes polyprotein

GAG_BAEVM
Baboon endogenous virus (strain M7)

Gag polyprotein (Core polyprotein)

GAG_FUJSV
Fujinami sarcoma virus

Gag polyprotein

GAG_GALV
Gibbon ape leukemia virus (GALV)

Gag polyprotein (Core polyprotein)

GAG_HV192
Human immunodeficiency virus type 1 group M subtype C (isolate 92BR025) (HIV-1)
GAG_HV193
Human immunodeficiency virus type 1 group M subtype F1 (isolate 93BR020) (HIV-1)
GAG_HV1A2
Human immunodeficiency virus type 1 group M subtype B (isolate ARV2/SF2) (HIV-1)
GAG_HV1AN
Human immunodeficiency virus type 1 group O (isolate ANT70) (HIV-1)
GAG_HV1B9
Human immunodeficiency virus type 1 group M subtype B (strain 89.6) (HIV-1)
GAG_HV1BR
Human immunodeficiency virus type 1 group M subtype B (isolate BRU/LAI) (HIV-1)
GAG_HV1ET
Human immunodeficiency virus type 1 group M subtype C (isolate ETH2220) (HIV-1)
GAG_HV1JR
Human immunodeficiency virus type 1 group M subtype B (isolate JRCSF) (HIV-1)
GAG_HV1Y2
Human immunodeficiency virus type 1 group M subtype B (isolate YU-2) (HIV-1)
GAG_HV1YB
Human immunodeficiency virus type 1 group N (isolate YBF106) (HIV-1)
GAG_HV2CA
Human immunodeficiency virus type 2 subtype A (isolate CAM2) (HIV-2)
GAG_HV2D1
Human immunodeficiency virus type 2 subtype A (isolate D194) (HIV-2)
GAG_HV2EH
Human immunodeficiency virus type 2 subtype B (isolate EHO) (HIV-2)
GAG_HV2G1
Human immunodeficiency virus type 2 subtype A (isolate Ghana-1) (HIV-2)
GAG_HV2KR
Human immunodeficiency virus type 2 subtype A (isolate KR) (HIV-2)
GAG_HV2RO
Human immunodeficiency virus type 2 subtype A (isolate ROD) (HIV-2)
GAG_HV2SB
Human immunodeficiency virus type 2 subtype A (isolate SBLISY) (HIV-2)
GAG_HV2ST
Human immunodeficiency virus type 2 subtype A (isolate ST) (HIV-2)
GAG_HV2UC
Human immunodeficiency virus type 2 subtype B (isolate UC1) (HIV-2)
GAG_JSRV
Sheep pulmonary adenomatosis virus (Jaagsiekte sheep retrovirus) (JSRV)

Gag polyprotein

GAG_KORV
Koala retrovirus (KoRV)

Gag polyprotein (Core polyprotein)

GAG_MLVAV
AKV murine leukemia virus (AKR (endogenous) murine leukemia virus)

Gag polyprotein (Core polyprotein)

GAG_MLVF5
Friend murine leukemia virus (isolate 57) (FrMLV)

Gag polyprotein (Core polyprotein)

GAG_MLVFF
Friend murine leukemia virus (isolate FB29) (FrMLV)

Gag polyprotein (Core polyprotein)

GAG_MLVFP
Friend murine leukemia virus (isolate PVC-211) (FrMLV)

Gag polyprotein (Core polyprotein)

GAG_MLVRD
Radiation murine leukemia virus

Gag polyprotein (Core polyprotein)

GAG_MPMV
Mason-Pfizer monkey virus (MPMV) (Simian Mason-Pfizer virus)

Gag polyprotein (Core polyprotein)

GAG_MSVMO
Moloney murine sarcoma virus (MoMSV)

Gag polyprotein (Core polyprotein)

GAG_SFV3L
Simian foamy virus type 3 (strain LK3) (SFVagm) (SFV-3)

Gag polyprotein (Pr71Gag)

GAG_SIVCZ
Simian immunodeficiency virus (isolate CPZ GAB1) (SIV-cpz) (Chimpanzee immunodeficiency virus)
GAG_SIVM1
Simian immunodeficiency virus (isolate Mm142-83) (SIV-mac) (Simian immunodeficiency virus rhesus monkey)
GAG_SIVS4
Simian immunodeficiency virus (isolate F236/smH4) (SIV-sm) (Simian immunodeficiency virus sooty mangabey monkey)
GAG_SIVSP
Simian immunodeficiency virus (isolate PBj14/BCL-3) (SIV-sm) (Simian immunodeficiency virus sooty mangabey monkey)
GAG_SMRVH
Squirrel monkey retrovirus (SMRV-H) (SMRV-HLB)

Gag polyprotein

GAG_SRV1
Simian retrovirus SRV-1

Gag polyprotein (Core polyprotein)

GAG_SRV2
Simian retrovirus SRV-2

Gag polyprotein (Core polyprotein)

GAG_WMSV
Woolly monkey sarcoma virus (WMSV) (Smian sarcoma-associated virus)

Gag polyprotein (Core polyprotein)

GAG_XMRV3
Xenotropic MuLV-related virus (isolate VP35) (XMRV)

Gag polyprotein (Pr65gag) (Core polyprotein)

GAG_XMRV4
Xenotropic MuLV-related virus (isolate VP42) (XMRV)

Gag polyprotein (Pr65gag) (Core polyprotein)

1 entry

P47(GAG-CRK) protein

GAGC_AVISC
Avian sarcoma virus (strain CT10)

37 entries

Matrix protein (Phosphoprotein M2)

Select_allDeselect_all
MATRX_ABLVB
Australian bat lyssavirus (isolate Bat/AUS/1996) (ABLV) reference strain
MATRX_CHAV
Chandipura virus (strain I653514) (CHPV) reference strain

Matrix protein

MATRX_DMSVA
Drosophila melanogaster sigma virus (isolate Drosophila/USA/AP30/2005) (DMelSV) reference strain

Matrix protein

MATRX_EBLV1
European bat lyssavirus 1 (strain Bat/Germany/RV9/1968) (EBLV1) reference strain

Matrix protein

MATRX_EBLV2
European bat lyssavirus 2 (strain Human/Scotland/RV1333/2002) (EBLV2) reference strain

Matrix protein

MATRX_MOKV
Mokola virus (MOKV) reference strain
MATRX_NDVB1
Newcastle disease virus (strain Chicken/United States/B1/48) (NDV) reference strain

Matrix protein

MATRX_PIV5
Parainfluenza virus 5 (strain W3) (PIV5) (Simian virus 5) reference strain

Matrix protein (Membrane protein)

MATRX_RABVP
Rabies virus (strain Pasteur vaccins / PV) (RABV) reference strain
MATRX_SENDO
Sendai virus (strain Ohita) (SeV) reference strain

Matrix protein (M protein)

MATRX_SENDZ
Sendai virus (strain Z) (SeV) (Sendai virus (strain HVJ)) reference strain

Matrix protein (M protein)

MATRX_TIBVC
Tibrogargan virus (strain CS132) (TIBV) reference strain

Matrix protein

MATRX_TUPVT
Tupaia virus (isolate Tupaia/Thailand/-/1986) (TUPV) reference strain

Matrix protein

MATRX_VSIVA
Vesicular stomatitis Indiana virus (strain San Juan) (VSIV) reference strain

Matrix protein

MATRX_VSNJO
Vesicular stomatitis New Jersey virus (strain Ogden subtype Concan) (VSNJV) reference strain

Matrix protein

MATRX_ABLVH
Australian bat lyssavirus (isolate Human/AUS/1998) (ABLV)
MATRX_ARAV
Aravan virus (ARAV)
MATRX_IRKV
Irkut virus (IRKV)
MATRX_ISFV
Isfahan virus (ISFV)

Matrix protein

MATRX_KHUV
Khujand virus (KHUV)
MATRX_RABVA
Rabies virus (strain PM1503/AVO1) (RABV)
MATRX_RABVB
Rabies virus (strain silver-haired bat-associated) (RABV) (SHBRV)
MATRX_RABVD
Rabies virus (strain China/DRV) (RABV)
MATRX_RABVE
Rabies virus (strain ERA) (RABV)
MATRX_RABVH
Rabies virus (strain HEP-Flury) (RABV)
MATRX_RABVI
Rabies virus (strain India) (RABV)
MATRX_RABVN
Rabies virus (strain Nishigahara RCEH) (RABV)
MATRX_RABVR
Rabies virus (strain China/MRV) (RABV)
MATRX_RABVS
Rabies virus (strain SAD B19) (RABV)
MATRX_SENDA
Sendai virus (strain Hamamatsu) (SeV)

Matrix protein (M protein)

MATRX_SENDF
Sendai virus (strain Fushimi) (SeV)

Matrix protein (M protein)

MATRX_SVCV
Spring viremia of carp virus (Rhabdovirus carpia)

Matrix protein

MATRX_VSIVC
Vesicular stomatitis Indiana virus (strain 94GUB Central America) (VSIV)

Matrix protein

MATRX_VSIVG
Vesicular stomatitis Indiana virus (strain Glasgow) (VSIV)

Matrix protein

MATRX_VSIVN
Vesicular stomatitis Indiana virus (strain 98COE North America) (VSIV)

Matrix protein

MATRX_VSIVS
Vesicular stomatitis Indiana virus (strain 85CLB South America) (VSIV)

Matrix protein

MATRX_WCBV
West Caucasian bat virus (WCBV)

5 entries

Nucleoprotein (Nucleocapsid protein)

Select_allDeselect_all
NCAP_MABVM
Lake Victoria marburgvirus (strain Musoke-80) (MARV) (Marburg virus (strain Kenya/Musoke/1980)) reference strain

Nucleoprotein (Nucleocapsid protein) (Protein N)

NCAP_MABVA
Lake Victoria marburgvirus (strain Angola/2005) (MARV)
NCAP_MABVO
Lake Victoria marburgvirus (strain Ozolin-75) (MARV) (Marburg virus (strain South Africa/Ozolin/1975))
NCAP_MABVP
Lake Victoria marburgvirus (strain Popp-67) (MARV) (Marburg virus (strain West Germany/Popp/1967))

Nucleoprotein (Nucleocapsid protein) (Protein N)

NCAP_MABVR
Lake Victoria marburgvirus (strain Ravn-87) (MARV) (Marburg virus (strain Kenya/Ravn/1987))

10 entries

Matrix protein VP40 (Membrane-associated protein VP40)

Select_allDeselect_all
VP40_EBORR
Reston ebolavirus (strain Reston-89) (REBOV) (Reston Ebola virus) reference strain
VP40_EBOZM
Zaire ebolavirus (strain Mayinga-76) (ZEBOV) (Zaire Ebola virus) reference strain
VP40_MABVM
Lake Victoria marburgvirus (strain Musoke-80) (MARV) (Marburg virus (strain Kenya/Musoke/1980)) reference strain
VP40_EBORE
Reston ebolavirus (strain Philippines-96) (REBOV) (Reston Ebola virus)
VP40_EBOSU
Sudan ebolavirus (strain Human/Uganda/Gulu/2000) (SEBOV) (Sudan Ebola virus)
VP40_EBOZ5
Zaire ebolavirus (strain Kikwit-95) (ZEBOV) (Zaire Ebola virus)
VP40_MABVA
Lake Victoria marburgvirus (strain Angola/2005) (MARV)
VP40_MABVO
Lake Victoria marburgvirus (strain Ozolin-75) (MARV) (Marburg virus (strain South Africa/Ozolin/1975))
VP40_MABVP
Lake Victoria marburgvirus (strain Popp-67) (MARV) (Marburg virus (strain West Germany/Popp/1967))
VP40_MABVR
Lake Victoria marburgvirus (strain Ravn-87) (MARV) (Marburg virus (strain Kenya/Ravn/1987))

19 entries

RING finger protein Z (Protein Z) (Zinc-binding protein)

Select_allDeselect_all
Z_LASSJ
Lassa virus (strain Mouse/Sierra Leone/Josiah/1976) (LASV) reference strain
Z_LATVB
Latino mammarenavirus (isolate Rat/Bolivia/MARU 1924/1965) (LATV) reference strain
Z_LYCVA
Lymphocytic choriomeningitis virus (strain Armstrong) (LCMV) reference strain
Z_PIRVV
Pirital mammarenavirus (isolate Rat/Venezuela/VAV-488/1995) (PIRV) reference strain
Z_TACVF
Tacaribe virus (strain Franze-Fernandez) (TCRV) reference strain
Z_TAMVU
Tamiami mammarenavirus (isolate Rat/United States/W 10777/1964) (TAMV) reference strain
Z_ALLVP
Allpahuayo mammarenavirus (isolate Rat/Peru/CLHP-2472/1997) (ALLV)
Z_BCNVU
Bear Canyon mammarenavirus (isolate Mouse/United States/AV A0070039/2000) (BCNV)
Z_CHAVB
Chapare mammarenavirus (isolate Human/Bolivia/810419/2003)
Z_CPXVB
Cupixi mammarenavirus (isolate Rat/Brasil/BeAn 119303/1970) (CPXV)
Z_GTOVV
Guanarito mammarenavirus (isolate Human/Venezuela/NH-95551/1990) (GTOV)
Z_IPPYV
Ippy mammarenavirus (isolate Rat/Central African Republic/Dak An B 188 d/1970) (IPPYV)
Z_JUNIN
Junin mammarenavirus (JUNV) (Junn mammarenavirus)
Z_MACHU
Machupo virus (MACV)
Z_MOBVC
Mobala mammarenavirus (isolate Rat/Central African Republic/Acar 3080/1983) (MOBV)
Z_OLVVA
Oliveros mammarenavirus (isolate Mouse/Argentina/RIID 3229/1990) (OLVV)
Z_PARVP
Parana mammarenavirus (isolate Rat/Paraguay/12056/1965) (PARV) (Paran mammarenavirus)
Z_SABVB
Sabia mammarenavirus (isolate Human/Brasil/SPH114202/1990) (SABV) (Sabi mammarenavirus)
Z_WWAVU
Whitewater Arroyo mammarenavirus (isolate Rat/United States/AV 9310135/1995) (WWAV)