Table 1 Regulatory factors of exosome biogenesis

HRS [30, 31]

STAM1 [32]

Positive correlated in various cells

HRS and STAM1 are members of ESCRT-0 complex which can recognize mono-ubiquitylated cargo proteins

TSG101 [33]

Positive correlated in various cells

TSG101 is a member of ESCRT-I complex and responsible for the budding of the endosomal membrane

CHMP4 [34]

Positive correlated in various cells

CHMP4 is member of ESCRT­III complex and perform fission of microdomains on the limiting membrane of MVEs, promoting vesicles separation

VPS4 [34]

Inhibition of VPS4b in HeLa cells increased exosome formation. Inhibition of both VPS4a and VPS4b in MCF-7 cells increased exosome formation, while inhibition one of VPS4a and VPS4b had no significant influence. Inhibition of both VPS4a and VPS4b in RPE1 cells inhibited exosome secretion

VASP4 plays an essential role in dissociation and recycling of the ESCRT complexes

ALIX [34]

Positive correlated in various cells

ALIX promotes endosome budding to form ILVs

nSMase2 [35]

Positive correlated in various cells

nSMase2 hydrolyzes sphingolipid to produce ceramide and promote MVBs budding inward

Cholesterol [36]

Positive correlated in oligodendroglia cell

They can sort related ligands into ILVs and sequestrate cytosolic proteins into ILVs

PLD2 [37, 38]

Positive correlated in MCF-7 cells

CD63 [39]

Positive correlated in MNT-1 melanoma cells

CD9 [40], CD82 [40]

Positive correlated in HEK293 cells

CD81 [41]

Positive correlated in human primary lymphoblasts

TSPAN8 [42]

Positive correlated in rat pancreatic adenocarcinoma cells

HSP70 [43, 44]

Positive correlated in various cells