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. 2021 Mar 1;40(7):e106922. doi: 10.15252/embj.2020106922

Table 1.

Proteins involved in membrane sealing. See text for references.

General function in membrane sealing Protein name Specific function Site of action
Sensing of membrane integrity Sensing of Ca2+ influx into cytosol PDCD6 Binds to ALIX to promote ESCRT recruitment Plasma membrane, endolysosomes
ALIX Promotes ESCRT recruitment Plasma membrane, endolysosomes
Annexins Together with Ca2+ binding proteins of the S100 family, involved in membrane fusion, shaping, sealing and tension reduction Plasma membrane, endolysosomes
SYT7 Endolysosomal protein that activates SNAREs in membrane fusion Plasma membrane
Dysferlin Accumulates phosphatidylserine at the site of membrane damage, as an “eat‐me” signal for macrophages. Interacts with some annexins Plasma membrane
TMEM16F Lipid scramblase that causes exposure of phosphatidylserine at the extracytosolic leaflet Plasma membrane
Sensing of β‐galactoside sugars on membrane proteins not normally exposed to cytosol GAL3 Interacts with ALIX and the autophagy‐associated ubiquitin E3 ligase TRIM16 Endolysosomes
GAL8 Inhibits mTORC1 and interacts with the autophagy receptor NDP52 Endolysosomes
GAL9 Activates AMPK Endolysosomes of macrophages and hematopoietic cells
Sensing of nuclear envelope integrity CHMP7 Binds to the inner nuclear membrane proteins LEMD2 and LEMD3 and recruits ESCRT‐III proteins Nuclear envelope, micronuclear envelope
Membrane fusion SNARE proteins (VAMP, syntaxin and SNAP families) Mediate fusion of specific membranes through formation of tetrahelical trans‐SNARE complexes Multiple
Membrane scission towards cytosol Acidic sphingomyelinase Lysosomal enzyme that is shed by Ca2+‐dependent exocytosis in response to wounding. Converts sphingomyelin into ceramide, which triggers caveolae endocytosis Plasma membrane
GRAF1 GTPase‐activating protein for the small GTPase Cdc42. Promotes clathrin‐independent endocytosis Plasma membrane, endosomes
Membrane scission away from cytosol ESCRT‐I Recruits ESCRT‐III Multiple
ESCRT‐III Filamentous complex that mediates scission of narrow membrane necks filled with cytosol, to achieve membrane sealing Multiple
VPS4 AAA ATPase that regulates ESCRT‐III assembly and disassembly Multiple